Flora of Peru

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" FLORA OF PERU

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CHARLES BAEHNI

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LUCIANO BERNARD:

CONSERVATOIRE ET JARDINS BOTANIQUES, GENEVA

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FIELD MUSEUM OF NATURAL HISTORY

VOLUME XIII, PART V-A, NUMBER 3
FEBRUARY 27, 1970

PUBLICATION 1087

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FLORA OF PERU

BY

CHARLES BAEHNI

AND

LUCIANO BERNARDI

CONSERVATOIRE ET JARDINS BOTANIQUES, GENEVA

BOTANICAL SERIES

FIELD MUSEUM OF NATURAL HISTORY

VOLUME XIII, PART V-A, NUMBER 3

FEBRUARY 27, 1970

PUBLICATION 1087

Library of Congress Catalog Card Number: 36-10^26

PRINTED IN THE UNITED STATES OF AMERICA
BY FIELD MUSEUM PRESS

FLORA OF PERU

SAPOTACEAE. Chicle family

REFERENCE: AubreVille, Sapotace'es, Me'moire I Adansonia 1964.
Baehni, Me"moire sur les Sapotace'es, III, Inventaire des genres, Bois-
siera 11, 1965.

Trees or shrubs, in rare instances (namely in some New Caledo-
nian species) undershrubs; pilosity composed of 2-armed hairs. Leaves
alternate, entire, in very rare cases opposite or subopposite, generally
with precociously caducous stipules, numerous parallel veins ("Ca-
lophyllum" blades) in several species of distinct genera (e.g., Chry-
sophyllum, Manilkara, Pouteria, etc.) or fewer distant veins. Flowers
small (except Tsebona, from Madagascar, when open, the calyx is
7.5 cm. in diameter), pedicels of variable length calyx with 4-5 or
8 sepals, frequently quincuncial or (Manilkara) in two valvate ver-
ticils (3 x 2), sepals free or partially connate (not more than the
half of the entire length of the calyx) , glabrous or silky inside, mostly
hirsute outside; corolla gamopetalous, tubular or broad campanulate,
with imbricate lobes (rarely quincuncial) without appendages (except
for Manilkara, dorsal appendages, and Bumelia with lateral ones),
staminodes present or absent, stamens (in the Peruvian genera) as
many as petals, filaments attached generally on the throat, ovary
mostly 5-celled, 1 ovule in each cell; fruit a berry with 1-5 seeds;
seed glossy with a basi-lateral or long ventral scar. Laticiferous ves-
sels always present.

The species are usually readily recognizable, but the separation of
the genera proves to be a difficult task, since, in all the family the
floral morphology is fairly uniform.

In the present case of the Peruvian flora, we would like to repeat
the introductory words of Prof. Mathias and Prof, Constance for the

135

136 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

family Umbelliferae (Flora of Peru, Field Mus. Bot., vol. XIII,
part V-A, no. 1, p. 4, 1962): "It is hoped that this preliminary re-
vision . . . will encourage collectors and Peruvian students of the
family"; our knowledge of the Sapotaceae of this country is inade-
quate. For instance, of the 39 species and one variety described
hereunder, 13 are known only from the Peruvian type; and 23 taxa
here presented, have no fruit or seed. It is universally admitted that
the taxonomy of this family is almost impossible without — for the
same species — flowers, fruit and seeds.

We trust that the reader will bear in mind the inadequacy of the
material available for this study and forgive us the imperfection to
be found in this work.

Since in the Sapotaceae the floral morphology is never sufficient for
the assignation to one or another genus, and because the form of the
seed scar is retained as a discriminative generic character, we think
it useful to include in the key for Pouteria, based on easy characters
of leaves, pedicels and flowers, the species: Mastichodendron Wil-
liamsii, Richardella glomerata var. glomerata and var. glabrescens and
Richardella macrophylla. For the same reason, in the key for Chry-
sophyllum, I added the Peruvian species of Cynodendron, separated
from the aforesaid genus on the basis of the seed morphology (Cyno-
dendron aurdtum, ferrugineum, ovale, revolutum), and also Gymnoluma
glabrescens, Prieurella Prieurii, Prieurella Wurdackii, their floral mor-
phology being very alike to Chrysophyllum.

I hope in this manner that the determination of samples devoid
of fruit and seeds, should still be possible; for the genera with few
species in Peru, as Achras (Calocarpum) , Bumelia, Manilkara, Sar-
caulus, which present easy and discriminating floral characters, the
generic key is sufficient.

Flowers 4-5 merous (in some instances 6-merous) with staminodes as
numerous as the stamens, fairly and uniformly developed.

Pouteria — and related taxa.

Flowers always 5-merous, without staminodes or with obsolete stami-
nodes, never as numerous as the stamens.

Chrysophyllum — and related taxa.

Calyx composed of 8 sepals, spirally disposed.

Achras (Calocarpum).

Calyx composed of 6 sepals on two verticils; corolla with append-
ages Manilkara.

FLORA OF PERU 137

Calyx composed of 5 valvate sepals; corolla without appendages.

Pouteria Bouffardiana.

Calyx composed of (usually) 4-5 quincuncial (rarely imbricate)
sepals.

Staminodes always present, equal in number to the stamens.

Lateral appendages of the corolla present Bumelia.

Lateral appendages absent.

Corolla fleshy, very thick, apparently valvate. Sarcaulus.

Corolla generally membranous, never very thick, imbricate

or quincuncial.

Ovules attached to the base of the loculi; seeds with a
basi-lateral scar and erect embryo.

Mastichodendron .

Ovules not attached to the base of the loculi.

Seed-scar somewhat narrow, more or less elongated.

Pouteria.

Seed-scar covering nearly the entire seed . . Richardella.

Staminodes absent or obsolete, irregularly present (1-2 per
flower) never in the same number as the stamens.

Inflorescences cauliferous (seed-scar lateral and narrow).

Prieurella.
Inflorescences ramiflorous.

Seed-scar lateral and narrow, seed 2 or more per fruit.

Chrysophyllum.
Seed, generally 1 per fruit.

Seed-scar lateral, broad and long Gymnoluma.

Seed-scar basilar or basi-lateral, very short. .Cynodendron.

ACHRAS L.

Calocarpum Pierre in Urb. Symb. Ant. 5: 97. 1904. Urbanella
Pierre, Not. Bot. Sapot. 25. 1890. Achradelpha 0. F. Cook, Journ.
Wash. Acad. Sci. 3: 160. 1913; Contrib. U. S. Nat. Herb. 16: 277.
1913.

REFERENCES: Aubre"ville, Adansonia 5: 16. 1965, PI. 1, p. 18 [to
read Achras sapota L. and not Nispero Achras (L.) Aubr., cf. Adan-
sonia 5: after p. 580. 1965] ; Gilly, Trop. Woods 73: pp. 5 and 20. 1945;
Fosberg, Taxon 13: 254. 1964.

138 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

A small genus containing perhaps 5-6 species characterized by the
calyx which is composed of 7-8 spirally arranged sepals, the outer
ones being smaller than the inner ones. Otherwise the flowers re-
semble those of the genus Pouteria. The seeds, as far as we know,
have a long and narrow scar.

The best known species, cultivated to some extent in Tropical
America is Achras zapota L. [sensu Aubre"ville, non auct.=Calocar-
pum mammosum Pierre] whose large ovoid fruits are edible.

Achras buchananiifolia (Pierre) Bernardi, Candollea 22: 230.
1967. Calocarpum buchananifolium (Pierre) Dubard, Ann. Mus. Col.
Marseille 20: 9. 1912. Urbanella buchananiaefolia Pierre, Not. bot.
Sap. 25. 1890.

Tree (?), twigs glabrous, slender, ribbed. Leaves membraneous,
narrowly oblong obovate, long acuminate at the tip, tapering to the
cuneate base in the young leaves, or with one acute base in the old
ones, entirely glabrous, 10-16 cm. long, 2-4 cm. wide; petioles gla-
brous, 8-11 mm. long, slightly canaliculate; midrib very prominent
on both sides, veins in 8-12 pairs, arcuate-ascendent, slender, obso-
lete above, more robust and visible below; venation forming a dense
and fairly conspicuous network; petioles subcanaliculate, 8-10 mm.
long. Flowers few, in the axils of leaves or above the scars, sessile,
near to 4 mm. long; sepals 8, the three external small (1-2 mm.),
triangular, hairy outside, the other 5 (3.5 mm. long) broad ovate,
thinner, ciliolate at the margins, glabrous outside, all the sepals, how-
ever, silky inside; corolla tubular, glabrous, 3 mm. long, lobes 5 of the
same length as the tube, broad-ovate, staminodes petaloid, fleshy,
oblong, 1 mm. long, fertile stamens with filaments very short, at-
tached to, or somewhat below the throat, anthers emarginate, less
than 1 mm. high, pistil 3 mm. high, with ovary globular, scarcely
1 mm. high, covered by long and erect silky hairs which hide the base
of the glabrous, stout style; stigma obsolete; ovary 5-celled, with
ovules narrowly attached from the inferior half of their adaxial side
to the basal area of the loculi. Fruit unknown.

Known only from the type. San Martin: near Tarapoto, Spruce

45U (G).

BUMELIA Sw.

REFERENCES: Candolle A. Prodr. 8: 154. 1844; Engler, Bot.
Jahrb. 12: 519. 1890; Baillon, Histoire des plantes 11: 277. 1891-
1892; Dubard, C. R. Acad. Sci. p. 393, feVr. 1911; Baehni, Candollea

FLORA OF PERU 139

7: 424. 1938; Record, Trop. Woods 59: 29. 1939; Brown & Clark,
Am. Journ. Bot. 27: 237. 1940; Clark, Ann. Mo. Bot. Card. 29: 155.
1942; Cronquist, Journ. Arn. Arb. 26: 445. 1945; Lloydia 9: 242.
1946; Bull. Torr. Bot. Club 73: 465. 1946; Meyer, Lilloa 13: 109.
1947; Wood & Channell, Journ. Arn. Arb. 41: 7. 1960; Chambers,
Trop. Woods 112: 40. 1960; Baehni, Boissiera 11: 134. 1965.

Shrubs or trees, often spiny. Inflorescences simple, ramiflorous;
flowers generally pentamerous, corolla lobes with a pair of lateral
appendages at the base. Berry 1-seeded (rarely 2-3), small; seed-
scar basilateral or basal; embryo vertical.

Bumelia obtusifolia Roem. & Schult., Syst. Veg. 4: 802. 1819;
Cronquist, Journ. Arn. Arb. 26: 456. 1945.

Shrub or small tree, often spiny. Leaves oblanceolate to broadly
ovate, rounded at the tip, initially sericeous beneath, later glabrate,
2-5 (-9) cm. long, 1-3 (-4.5) cm. wide. Flowers solitary or numer-
ous in axillary clusters, subsessile or in pedicels up to 8 mm. long;
sepals 1.3-3 mm. long, strigose or sericeous; free lobes of the corolla 5,
as long as or longer than the tube, each with a pair of lateral append-
ages; staminodes from narrowly oblong and erose to ovate and sub-
entire, sometimes hooded, about 1.5-3.2 mm. long; style about 1.5-
4.6 mm. long. Berry small, ellipsoid-cylindric to subglobose, about
8-15 mm. long, seed obovoid, scar circular, "glabellae" about half
as high as the seed.

Cajamarca: "in ripa fluminis Amazonum, ad confluentem Chin-
chipen," Humboldt & Bonpland.

CHRYSOPHYLLUM L.

REFERENCES: Cronquist, Bull. Torr. Bot. Club 73: 287. 1946;
AubreVille, Adansonia 1: 9. 1961; Baehni, Boissiera 11: 71. 1965.

Trees or shrubs; under surface of leaves often sericeous, the hairs
malpighiaceous, or stalked and bifurcate; venation of various types.
Flowers in axillary clusters, generally 5-merous (rarely 4-6 merous),
sympetalous; staminodes wanting, or rarely a few casual and irreg-
ularly developed vestiges; ovary (2)-5-(10)-celled; ovules attached
laterally or basilaterally to the axial placenta; style short columnar.
Fruit generally fleshy, indehiscent, with (1-) 4-5 seeds, rarely more,
with narrow or moderately broad basilateral scar.

Adult leaves glabrous or with a few scattered hairs beneath.

140 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

Flowers pedicellate; pedicels twice (or more) as long as the flowers,
corolla glabrous.

Leaf blades membranous, elliptical, gradually acuminate, ter-
tiary veins almost obsolete ; fascicles of flowers on the young
branchlets Cynodendron ovale.

Leaf blades coriaceous obovate; fascicles generally on the old,
leafless branch.

Venation network fairly conspicuous on both sides; blades
more than 10 cm. long Prieurella Wurdackii.

Venation network obsolete; blades less than 8 cm. long.

Chrysophyllum Wurdackii.
Flowers almost sessile, or with pedicels as long as the flowers.

Corolla densely hairy, rust-coloured hairs.

Cynodendron revolutum.
Corolla glabrous.

Corolla lobes as long as the tube, filaments inserted on the
throat; membranous blades with secondary veins and
fairly distinguishable network . . Chrysophyllum albipilum.

Corolla lobes 3 times longer than the tube; filaments inserted
at the base of the tube; fleshy-coriaceous blades with
quite obsolete network Gymnoluma glabrescens

Adult leaves covered beneath with white or coloured hairs.

Flowers pedicellate, twice (or more) longer than the flowers, gen-
erally on the old branches Prieurella Prieurii.

Flowers almost sessile, or with pedicels as long as the flowers.
Corolla hairy outside.

Calyx sericeous inside; blades generally oblong, persistently
rufous-sericeous below Cynodendron ferrugineum.

Calyx glabrous inside; blades generally elliptic, sometimes be-
coming glabrate with age Cynodendron auratum.

Corolla glabrous.

Stamens exserted ; blades less than 8 cm. long, petioles barely
1 cm. long Chrysophyllum granatense.

Stamens included; blades more than 15 cm. long, petioles
3-4 cm. long Chrysophyllum Ulei.

Chrysophyllum albipilum Cronq. Bull. Torr. Bot. Club 73:
299. 1946.

FLORA OF PERU 141

Small tree, 7 m. high (collected only once!), branchlets slender,
terete, sparsely white pilose. Leaves membranous, elliptic, at the bor-
der slightly undulate, dull on both sides, soon glabrate above, white-
shining sparse-minutely puberulous beneath, 4-11 cm. long, 2-4.5 cm.
wide, however frequently 6 x 2.5 cm.; midrib hardly impressed above,
scarcely prominent beneath; secondary veins about 7-10 pairs, obso-
lete on the upper side, slightly raised on the back, arcuate near the
margins; veins rather irregular, forming a very loose reticulum; peti-
oles 2-4 mm. long, hardly canaliculate, white appressed pilose. Flow-
ers pentamerous, by 2-4 in the axil of leaves, pedicels gray-sericeous,
2 mm. long; calyx quincimcial, deeply lobed, 2 mm. high, outer sepals
ovate-acute, the inner ones obtuse, sparsely pilose on the back, gla-
brous inside, ciliate and thinner at the margin, elsewhere fleshy;
corolla green, glabrous, 4 mm. high; lobes 5, ovate, quincuncial,
about half as long as the tube; filaments very short, nipple-shaped,
attached to the throat, anthers 0.5-0.6 mm. long with divergent cells;
pistil conical, scarcely 1.5 mm. high, with ovary depressed, finely
pilose, contracted to the thick, glabrous, finely 5-furrowed style,
1 mm. long, stigma lobes (4) 5. Ovary (4) 5-celled, with loculi pro-
portionally spacious, ovules attached near the base of the loculi.
Fruit unknown.

San Martin: Juan Jui, Alto Rio Huallaga, 400-800 m., King 4365,
type (F, G).

Chrysophyllum granatense Spreng. Syst. 1: 667. 1825. Nyc-
terisition argenteum H.B.K. Nov. Gen. & Sp. 3: 238, t. 244. 1818.

Tree (?) known only from the type! Branchlets terete, somewhat
wrinkled in the herbarium sample, appressed silky tomentose toward
the ends as also on the elongated leaf buds. Leaves elliptic (rarely
elliptic-obovate) , coriaceous, 4-5 (6) cm. long, 2-2.5 (3) cm. wide,
glabrous and shining above, white-sericeous beneath, midrib slender
and impressed above, fairly robust and prominent beneath; blades
closely and conspicuously striate above, the thread-like veins almost
parallel, averaging 0.5 mm. apart, jointed near the border, fairly
obsolete beneath; petioles 5 (8) mm. long, semiterete, canaliculate,
silky. Flowers 6-9 in axillary fascicles, slender and silky pedicels 3-4
mm. long; calyx quincuncial (in the unique sample at our disposal,
4 and 6-merous flowers were also observed) ; sepals almost completely
free, large-ovate, scarcely more than 1 mm. high, silky puberulous
on the back, glabrous inside; corolla glabrous inside, with some silky
hairs outside, fairly fleshy, with a very short tube, less than 1 mm.

142 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

long, petals ovate, reflexed, 1.5 mm. high; fertile stamens exserted
(4) 5 (6), with slender and subulate filaments attached at the top
of the tube, anther oblong-acute, cordate and bilobed at the base,
dorsifixed; pistil 3 mm. high, with a glabrous and slender style nearly
2 mm. long, stigma punctiform, almost obsolete; ovary tawny-hir-
sute, (4)- 5- (G)-locular, ovules attached by the middle of their
adaxial side.

Known only from the type collection. Cajamarca: near Jae*n (de
Bracamoros) 550-600 m. alt. Humboldt & Bonpland s/n (P).

Chrysophyllum Ulei Krause, Notizbl. Bot. Gart. Berlin 6: 171.
1914; Baehni, Boissiera 11: 74. 1965. Ecclinusa Ulei (Krause) Gilly
ex Cronq. Bull. Torr. Bot. Club 73: 311. 1946. Ragala Ulei (Krause)
AubreV. Adansonia 4: 368. 1964. Ecclinusa balata Ducke, Arch. Jard.
Bot. Rio de Janeiro 6: 76. 1933; Aubre"ville, I.e. Chrysophyllum ba-
lata (Ducke) Baehni, Boissiera 11: 75. 1965. Chrysophyllum Klugii
Baehni, Candollea 7: 134. 1936. Ecclinusa Klugii Cronq. Bull. Torr.
Bot. Club 73: 311. 1946.

Medium-sized or tall trees up to 40 m. high, branchlets robust,
strongly compressed at the end, almost square in transverse section,
rusty or ochre, short appressed pilosity, hairs, however, caducous with
age. Leaves coriaceous, large elliptical to obovate, frequently with
a large and short acumen, rounded at the base, 16-20 (30) cm. long,
9-11 cm. wide; blades on the upper side soon becoming glabrate,
covered below with glossy, appressed, reddish hairs, which, in some
cases fall more or less completely, the under surface glabrate and dull ;
petiole very strong, 3-4 (5) cm. long, canaliculate; midrib fairly im-
pressed above, strongly prominent beneath; secondary veins, 15-20
pairs, conspicuously raised beneath, parallel, straight except near the
margin, where they are slightly arched; tertiary veins almost obso-
lete not visible without a lens, dense, equidistant, more or less paral-
lel. Flowers fasciculate in the axils or above the leaf scars; pedicels
stout, short, ochre-colour or rusty tomentose, scarcely 2 mm. long;
sepals 5 (6 ?) ovate, obtuse, almost completely free, up to 3-4 mm.
high, quincuncial, hairy on the back like the pedicels, glabrous inside;
corolla glabrous, campanulate, 5 mm. high, tube only 1.5 mm. long,
lobes 5, imbricate, broadly ovate; fertile stamens 5, with stout fila-
ments, attached to the middle of the tube, 2 mm. long, anthers large,
1 mm. high, apiculate, basifixed; pistil 4 mm. high, rather conical,
ovary long pilose, approximately 2 mm. high, style attenuate, stigma
barely thickened; ovary 5 (6) -celled, locules opened just at the base

FLORA OF PERU 143

of the carpels, ovules attached by their basal-ventral side. Berry be-
coming almost ligneous with age, globular-depressed, at the beginning
covered with short tawny hairs, later glabrate, approximately 3 cm.
in diameter, supported by the accrescent, nearly lignified calyx ; seeds
2-3 (more ?), free, nearly 2 cm. long, with scar rather narrow and
short (after Ducke I.e.).

Type: Ule 8729 (B) destroyed in the past war; I do not know if
duplicates of this number are extant.

Guyana, Brazil (Amazon), Peru. Lore to: Tierra Doble on the
Rio Nanay, Williams 911 (F).- — San Martin: Zepelacio, near Moyo-
bamba, 1100 m., King 3699 (F).

To this species could belong two sterile specimens of LI. Williams,
from Loreto: 907 (Tierre Doble on the Rio Nanay) and 929 (Tim-
buchi on the Rio Nanay). However, because of the glabrate under-
surface of the blades, they resemble even more Chrysophyllum san-
guinolentum (Pierre) Baehni.

Chrysophyllum Wurdackii (Aubr.) Bernardi, Candollea 22:
231. 1967. Noxythece Wurdackii Aubr. Adansonia 5: 201. 1965.

Tree 25 m. high (only once collected) with stout, erect, greyish
branchlets, terete except for the compressed, furrowed and sparsely
rusty puberulent extremities. Leaves obovate, fleshy-coriaceous,
generally rounded or very obtuse at the tip, cuneate-elongate toward
the base, merging gradually into the flat and sparsely silky pilose
petiole only 5 mm. long; blades 4-6 cm. long, 1.5-2 cm. wide, some-
what wrinkled (in botanical samples), glabrous and opaque above,
whitish (glaucous when fresh) beneath with scattered small, silky
hairs; midrib flat on the upper surface, imperceptibly prominent on
the reverse side; lateral veins in 8-10 pairs, very thin and scarcely
noticeable on both sides; veinlets and network obsolete. Fascicles
10-15-flowered, in the axils of the leaves and leaf-scars and on first
year's newly lignified branchlets; pedicels very loosely puberulent,
4-5 mm. long, gradually swollen toward the receptacle; calyx quin-
cuncial, opening very early in the bud's development, composed of
sepals having their lower halves connate to the receptacle, with very
scattered reddish hairs, lobes very broad-ovate, 2 mm. wide, 1-1.25
mm. long, glabrous inside, fleshy, with nearly scarious margins; co-
rolla tubular, about 2 mm. high, fleshy, glabrous, with imbricate
petals broad-ovate, half as long; 5 stamens, 1 mm. high with very
short filaments attached at the throat, anthers broad, obtuse, basi-
fixed; staminodes generally absent, or only 1-2, very small and irreg-

144 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

ular; pistil broad, flatted conical, 2 mm. wide at the base, 1 mm. high,
pubescent, with a style half as long, evenly decreasing in diameter
up to the small, adnate stigmas; ovary bi-celled, with 2 ovules nar-
rowly attached by the middle of their adaxial side. Fruit unknown.

Known only from the type. Amazonas: Prov. Bagua, forested
ridge on right bank of Rio Santiago 2-3 km. above mouth, elev. 300-
350 m., Wurdack 2^09, Type (F).

CYNODENDRON Baehni

REFERENCES: Baehni, Arch. Sci. Geneve 18: 34. 1965; Boissiera
11: 142. 1965; AubreVille, Adansonia 6: 192. 1966.

An American genus of medium-sized trees having coriaceous
leaves, frequently silky beneath and with numerous parallel veins,
flowers in axillary fascicles, calyx 5-merous, imbricate or quincuncial,
5 (6-7) petals without appendages, 5 stamens, staminodes absent,
ovary (4) 5 (6-7)-locular, medium or small-sized fruit, generally
1-seeded (rarely 2 or more, free), seed-scar small or very small, lateral
or basilateral.

The species of Cynodendron were previously in the genus Chry-
sophyllum; for the floral characters and general vegetative appear-
ance, Cynodendron is very similar to that of the old Linnean genus;
however, since the taxonomy of the Sapotaceae is based, according to
all the recent authors, not only on the floral morphology, but also
on the seed scars, Cynodendron must be accepted.

For specimens devoid of fruit, the identification of Cynodendron
or Chrysophyllum species will always be hypothetical; only sound
floristic and phytogeographical studies could help the taxonomist in
this event.

Cynodendron auratum (Miq.) Baehni, Boissiera 11: 143. 1965.
Chrysophyllum auratum Miq. in Mart. Fl. Bras. 7: 27. 1863. Chry-
sophyllum auratum var. majus Miq. I.e. Chrysophyllum sericeum
A. DC. Prodr. 8: 158. 1844 (non Salisb. Prodr. p. 138. 1796. Chry-
sophyllum cainito L.).

Tree, small or medium-sized, up to 30-35 m., rarely a shrub;
branchlets fairly robust, flexuose, terete and ochre-yellow except the
ends which are compressed and rusty-sericeous, eventually becoming
glabrate. Leaves membranous, elliptic, ovate to obovate, rarely ob-
long, acuminate or abruptly acuminate at the tip, acute or subcune-
ate at the base, glabrous above, more or less persistently bronze-silky

FLORA OF PERU 145

beneath, in some instances almost glabrate, 10-15 cm. long (rarely
more or less) and 5-8 cm. wide; petioles appressed rusty pubescent,
fairly canaliculate, 10-15 (20) mm. long; midrib thin impressed
above, strong prominent beneath; lateral veins well-spaced and arch-
ing, 14-18 pairs, scarcely noticeable above, thin but distinctly raised
beneath, the other veins almost obsolete. Flowers 10-20 in the axil
of leaves or even more numerous above the leaf scars; rarely on short
lateral and defoliated twigs; pedicels 3-5 mm. long, bronze-silky
slender when elongated ; calyx pentamerous, very thick, lobes almost
free, quincuncial, thinner near the margin, large ovate, almost orbic-
ular, rusty hairy on the back, glabrous inside or, in rare instances,
with few hairs, 2 mm. high; corolla tubular, more or less densely
silky outside except at the upper end of the lobes, 4-5 mm.
(rarely more) long; lobes 5 (-6), ovate, rounded, 1-2 mm. long, im-
bricate or — observed only in Guyana specimens — quincucial; fila-
ments very short, stout, attached to the throat, anther obtuse, with
thecae divergent at the base; pistil almost conical, with a hirsute
ovary, 5 (6)-celled, ovules attached toward the base, style 1 (-2) mm.
long. Berry wide-ellipsoid, about 1.5-2 mm. long; seed solitary with
broad basilateral obcordate scar extending to about the middle.

Type: Schomburgh 864, Roraima, British Guiana.

Guianas, Venezuela, Colombia, Brazil, Peru. Loreto: Mishu-
yacu, near Iquitos, 100 m. alt.: Klug 1318 (F); Lower rio Huallaga,
155-210 m. alt.: Williams 3945 (F, S).

Cynodendron ferrugineum (Ruiz & Pav6n) Bernardi, Can-
dollea 22: 231. 1967. Nycterisition ferrugineum Ruiz & Pav6n, Fl.
Peruv. 2: 47, Ic. 187. 1794; Prodr. 30, Ic. 5. 1794. Chrysophyllum fer-
rugineum (Ruiz & Pav6n) Steud., Nomencl. ed. 2, 1: 359. 1840 [non
Gaertn. f. Fruct. 4, 122, tab 202, fig. 1, 1805. Cynodendron oliviforme
(L.) Baehni]. Chrysophyllum Pavonii Cronq. Bull. Torr. Bot. Club 73:
306. 1946.

Medium-sized tree, up to 25 m. high (ex Ruiz & Pa von, I.e.);
branchlets slender, light brown, densely rusty-red pilose at the ends,
compressed, becoming glabrate. Leaves elliptic to oblong, membra-
nous, acute to subacuminate at the tip, acute at the base, glabrous
above except at the base of the impressed midrib, rufo-sericeous be-
low, 8-12 cm. long, 3-5 cm. wide; petioles rusty-pilose slightly angu-
late and canaliculate, 8-10 mm. long; midrib beneath fairly promi-
nent; lateral veins numerous; more than 20 pairs, very thin and
raised on both sides, bifurcate near the margins. Flowers numerous

146 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

on short lateral defoliated pilose twigs, at the axil of leaves or above
the leaf scars, pedicels sericeous, compressed, 1-2 mm. long; calyx
pentamerous, 2 mm. high, with sepals ovate 1 mm. long, thick coria-
ceous, hairy outside, sericeous inside; corolla tubular, pentamerous
hairy outside, except at the end, up to 4 mm. long; petals imbricate,
nearly 2 mm. long, ovate; filaments very short, attached on the throat,
anthers small, obtuse; ovary globose, 1-1.5 mm. wide, hirsute, 5-
celled, loculi opened near the base, ovules attached at the middle of
their ventral side or toward the bottom; style very short, stigma with
5 small protuberances. Berry unknown.

Type: Ruiz "Nycterisition ferrugineum, verancule Chichimicuna"
(G).

Brazil (Rio Jurua, Sao Sebastiao, Ule 5164}- — Loreto: Iquitos,
shore of Rio Itaya, Asplund 11+278 (S)- — San Martin: Juan Jui, Alto
Rio Huallaga, 400 m. alt., King 3843 (F, S)— Huanuco: "Cuchero,
Chinchao et Pillao ad Chacahuassi [sic], nemoribus imis et calidis,"
Ruiz & Pavon (G, F, MA).

Cynodendron ovale (Rusby) Bernard!, Candollea22: 231. 1967.
Chrysophyllum ovale Rusby, Mem. N. Y. Bot. Gard. 7: 320. 1927;
Cronquist, Bull. Torr. Bot. Club 73: 304. 1946.

Medium-sized or tall tree up to 25-40 m. high; branchlets slender,
grey, coarsely angled at the end, appressed rusty pubescent, soon
glabrous. Leaves membranous, elliptic, less frequently elliptic-ovate,
abruptly acuminate, rounded or obtuse at the base, glabrous except
for a few scattered hairs below, 6-11 cm. long, 2.5-6 cm. wide, peti-
oles white pilose, becoming glabrate, scarcely canaliculate, 4-6 mm.
long; midrib impressed above, prominent but slender beneath; 15-
20 pairs of lateral nerves, very thin and slightly raised on both sides.
Flowers, 10-20 in the leaf axils or above the leaf scars; in some in-
stances, fascicled on a very short lateral leafless twig; pedicels slender,
white puberulent, 4-6 mm. long; calyx quincuncial with moderately
thick sepals with thinner borders, broadly ovate, almost completely
free, sparingly pilose on the back, sericeous inside, 1 mm. high; co-
rolla quincuncial, glabrous, fleshy, campanulate, 3 mm. long, lobes 5,
ovate, scarcely 2 mm. long; stamens less than 1 mm. long, with fila-
ments attached to the throat, small anthers with divergent cells;
pistil globular, 1 mm. diameter, finely tawny pilose with stigma al-
most sessile and with five small lobes; ovary 5-celled, with ovules
attached by the middle of their ventral side. Fruit "globose or a
little longer than broad, about 1.5 cm. broad, fleshy, mostly 1-seeded

FLORA OF PERU 147

in my specimens, the seed 1 cm. or more long and more than half as
broad, ellipsoid, light-brown, somewhat shining, the gray hilum tak-
ing up two-thirds of the length and half the circumference of the
seed." (After Rusby, I.e.)

Type: 0. E. White 1384-A (NY), Bolivia, Esperanza Falls (not
seen).

Brazil, Peru, Bolivia. — Loreto: Florida, Rio Putumayo, at mouth
of Rio Zubineta, alt. 180 m., King 2324 (F, G, S).— - Madre de Dios
(or: Brazil ?): Rio Acre, Monte M6, Ule 9694 (G).

Cynodendron revolutum (Mart. & Eichl.) Bernardi, Candollea
22: 231. 1967. Chrysophyllum revolution Mart. & Eichl. in Mart. Fl.
Bras. 7: 104. 1863.

Medium-sized tree, branchlets compressed at the extremity, al-
most 4-angular, ochre-yellow pubescent becoming glabrate. Leaves
elliptic, rarely elliptic-oblong, almost coriaceous, revolute at the mar-
gin, eventually glabrescent above except on the midrib, puberulent
beneath with forked hairs more dense along the midrib, 6-9 cm. long,
3-5 cm. wide, petioles stout, rufo-hirsute, semi-terete, 5 mm. long;
midrib stout and fairly prominent beneath, obsolete and slightly im-
pressed above; lateral veins about 10-15 pairs a little raised beneath,
nearly obsolete above, spreading and arched toward the margin.
Flowers sessile or sub-sessile, 8-10 in the axils of leaves or above the
leaf scars; pedicels, when present, very short, stout and rufo-hirsute;
calyx pentamerous, with almost completely free sepals, triangular,
nearly 2 mm. long, very thick, rufo-hirsute on the back, appressed
pilose inside; corolla tubular, fleshy, densely hairy outside except
below the sepals and at the tips of the lobes, glabrous inside, 4-5 mm.
long, lobes 3, quincuncial, ovate, nearly 1 mm. long, filaments very
short, attached to the throat, anthers 0.4-0.5 mm. long; pistil glo-
bose, nearly 2 mm. long, densely rufous hairy except at the end of the
short style (up to 0.5 mm. long);ovary 5-celled, loculi opened at the
middle, ovules attached by the central portion of their ventral side.
Berry ellipsoid-globose, about 12 mm. long, single-seeded; seed-scar
basilateral, very broad and long, more than rather half the length of
the seed.

Type: Spruce 4260, "prope Tarapoto, Peruvia orientalis."

San Martin: Spruce 4260 (F, G). Tarapoto, 750 mi. Williams
5522; 6140 (F); Alto Rio Huallaga, 360-900 m. Williams 6655 (F).
Juan Guerra, 720 m. Williams 6900 (F).

148 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

GYMNOLUMA Baill.

REFERENCES: Engler, in Engl. & Prantl, Nat. Pflanzenf. Nachtr.
p. 274. 1897; Aubre"ville, Adansonia 1: 26, t. 10, fig. 8. 1961; Baehni,
Boissiera 11: 100. 1965.

Trees; flowers fasciculate, calyx without appendices, pentamer-
ous, petals 5, almost free, staminodes completely lacking, or few
(1-2) and irregular, ovary (in American species) tri-locular, globose
fruit one-seeded with long and large scar, vertical embryo.

Gymnoluma glabrescens (Mart. & Eichl.) Baillon, Hist. PI. 11:
292. 1891. Lucuma glabrescens Mart. & Eichl. in Mart. Fl. Bras. 7:
72, t. 46, fig. 1. 1863. Vitellaria glabrescens (Mart. & Eichl.) Radlk.
Sitzb. Math.-Phys. Cl., Acad. Wiss. Miinchen 12: 326. 1882. Pouteria
glabrescens (Mart. & Eichl.) Baehni, Candollea 9: 350. 1942. Oxy-
thece Ferreirii Cronq. Bull. Torr. Bot. Club 73: 468. 1946. Elaeoluma
glabrescens (Mart. & Eichl.) Aubr. Adansonia 1: 26. 1961.

Tree, with stout branchlets, almost cylindrical in the lower por-
tion of the samples, slightly compressed at the ends, light-brown, dull
and glabrous. Leaves alternate, at the ends of the branchlets, becom-
ing sub-opposite, fleshy-coriaceous, glabrous, dull on both sides, very
peculiar in herbarium samples for the inferior side ochre-coloured;
elliptical or oblong, 8-15 (20) cm. long, 5-7.5 cm. wide, petioles
10-15 mm. long, stout and flat on the upper side; midrib prominulus
beneath; secondary veins in 10 or more pairs, very slender, patent
and slightly arcuate near the edges of the blade, as a rule, never paral-
lel. Flowers 5-10 in the axils of the leaves, or above the leaf scars,
pedicels glabrous and robust, 3-5 mm. long; calyx pentamerous, quin-
cuncial, the outer sepals broadly ovate and coriaceous, the two inner
thinner and elliptical, 4-6 mm. long; corolla 5-6 mm. long, membra-
naceous with tube very short, petals 5, elliptical, 4 mm. long; stamens
5, filaments robust, 1 mm. long, attached near the base of the corolla,
anthers dorsifixed, sagittate, 2 mm. long; staminodes not observed;
pistil 2 mm. long, with ovary nearly cylindrical and half as long as
the whole pistil, minutely tomentose, style stout and glabrous, with
obtuse and inconspicuous stigma; loculi 3 (5), with the ovules at-
tached at the top. Berry globose, smooth, 2 cm. diameter, one-
seeded; seed ellipsoid 10 x 8 mm., with a long and large scar, con-
spicuously warty.

Type: Spruce 2029, Rio Negro, between Barcellos and S. Gabriel,
Brazil.

FLORA OF PERU 149

Amazonian Brazil and Peru (Venezuela, Colombia ?). Dept.
Lore to: Iquitos and vicinity; Williams 3669 (F).

MANILKARA Adans.

REFERENCES: Reede, Hort. Malab. 4: t. 25. 1683; Dubard, Ann.
Mus. Col. Marseille 23: 6. 1915; Cronquist, Bull. Torr. Bot. Club 72:
550. 1945; Monachino, Phytol. 4: 94. 1952; Baehni, Boissiera 11:
91. 1965.

Trees frequently tall, or sometimes shrubby, with hard red wood ;
inflorescences simple, ramiflorous, fasciculate, flowers usually with
long and slender pedicels; sepals 3+3, petals 6, thin, each with two
dorsal appendages; stamens as many as the corolla-lobes, alternating
with an equal number of staminodes; ovary 6-14-loculate; berry
1-2 (4)-seeded, with seeds free, seed-scar narrow and short, embryo
upright.

Manilkara surinamensis (Miq.) Dubard, Ann. Mus. Col. Mar-
seille, 23: 22. 1915. Mimusops surinamensis Miq. in Mart. Fl. Bras.
7: 43. 1863. Mimusops amazonica Huber, Bol. Mus. Goeldi 4: 433.
1904. Mimusops maparajuba Huber, I.e. p. 434. Manilkara ama-
zonica (Huber) Chev. Rev. Bot. Appl. 12: 276. 1932. Manilkara
bidentata Williams [non (A.DC.) A. Chev.], Field Mus. Publ. Bot.
15:412. 1936.

Medium-sized tree, up to 30 m. high; branchlets thick, light
brown, glabrous, covered with leaf scars. Leaves papery, rarely cori-
aceous, slightly obovate to elliptical, rounded or slightly emarginate,
in some instances subacute at the tip, cuneate at the base, glabrous
on both sides, with a strongly prominent midrib beneath, secondary
veins very numerous, thin and parallel, tertiary ones forming an ob-
solete but dense network; blades 8-12 cm. long, 3-5 cm. broad; peti-
oles slender, glabrous, canaliculate, 15-30 mm. long. Flowers whitish,
generally crowded at the ends of the branchlets, in few-flowered (3-5)
but numerous fascicles; slender and glabrous pedicels (10) -12- (16)
mm. long; the 3 outer sepals glabrous or covered with deciduous
squamulae, the 3 internal ones whitish pubescent, 3-3.5 x 2 mm.;
corolla appendages narrow-ovate, 3.5 mm. high, 1-1.2 mm. wide,
petals alike, concave and connate at the base, forming a tube 1 mm.
high; 6 fertile stamens 3-3.2 mm. long, with stout dorsifixed anthers
and filaments 2 mm. long; staminodes as long as the stamens, 2-4
-dentate or -laciniate, with divisions generally less than 1 mm. long;
pistil 5-5.5 mm. high, with more or less conical ovary quite glabrous

150 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

less than 2 mm. high, style stout, attenuated toward the obscurely
bi-lobed stigma; ovary 6-7-celled. Berry turbinate-globose, 15-18
mm. wide, with 1-2 seeds compressed, up to 15 mm. long.

Type: Spruce 3351 "Habitat ad flumina Cassiquiari, Vasiva et
Pacimoni" (Venezuelan Amazonia).

Venezuela, Brazil, Peru. Loreto: Tierra doble on the Rio Nanay,
Williams 893 (F, G) ; Caballo-Cocha, on the Amazon River, Williams
2260 (F) ; Lake Victoria, on the Amazon River, Williams 2572a (F)—
S. Martin: Alto Rio Huallaga, 360-900 m. Williams 5735 (F)— Hua-
nuco: Prov. Pachitea, Distr. Honoria, mouth of the Rio Pachitea,
Raphael Lao M. 56 (fl.), 90 (fr.). "Quinilla, quinilla colorada."

Chrysophyllum Michino H.B.K. Nov. Gen. et Sp. 3: 236. 1818 de-
scribed on sterile material, and never more discovered, very probably
pertains to this species ("Provinciae Bracamorensis, prope Cavico et
Matara") Humboldt & Bonpland 3589.

MASTICHODENDRON Cronq.

REFERENCES: Lam, Rec. Trav. Bot. Ne"erl. 36: 521. 1939; Dug-
and, Caldasia 4: 427. 1947; van Royen, Blumea 10: 122. 1960;
AubreVille, Adansonia 3: 32. 1963; Baehni, Boissiera 11: 126. 1965.

Sideroxylon sect. Mastichodendron, Engler, Bot. Jahrb. 12: 496.
1890; id. in Engler & Prantl, Pflanzenf. 4, 1: 144. 1891.

Trees; inflorescences simple — sometimes apparently branched—
ramiflorous ; flowers pentamerous, calyx without appendices, corolla
bell-shaped with a short tube, staminodes present ; fruit indehiscent,
1-seeded (rarely 2-3, free), scar short, basilar or basi-lateral, embryo
vertical.

Mastichodendron Williamsii (Baehni) Baehni ex Bernardi,
Candollea 22: 231. 1967. Sideroxylon Williamsii Baehni, Candollea
7: 135. 1936.

Tree 20 m., with large crown, and diameter 50-75 cm.; branchlets
slender grisaceous, puberulous at the tip. Leaves coriaceous, elliptic,
obtusely cuspidate (acumen up to 5 mm. at maximum), acute at the
base, glabrous above, sparsely pilose beneath ; secondary veins very
numerous and parallel, very thin above, quite obsolete beneath ; 5-
8 cm. long, 2.5-3 cm. wide, petioles 6-10 mm. Flowers (seen only
in a young stage) 3-5 in the axils of the persistent leaves; pedicels
3 mm. long; calyx pentamerous, quincuncial, sepals orbicular, rufous-
pubescent on the back, silky inside; corolla pentamerous, glabrous,

FLORA OF PERU 151

with tube 1 mm. long and lobes 1.5 mm. high; stamens 5, filaments
very short, attached to the throat; staminodes stipitiforme, on the
same level, truncate; ovary rufous-hirsute, 5-celled, ovules attached
to the lower end ; style glabrous, as long as the oxary, black ; stigma
entire, punctiform. Berry not seen (after LI. Williams: yellowish at
maturity) .

Known only from the type. Loreto: Palta Cocha on the upper
Rio Nanay, Williams 3198 (F: type).

POUTERIA Aubl.

REFERENCES: A. De Candolle, Prodr. 8: 165. 1844; Bentham &
Hooker, Genera Plant. 1: 658. 1876; Engler, Bot. Jahrb. 12: 514.
1890, in Engl. & Prantl, Nat. Pflanzenfam. 4, 1: 141. 1891; Kuntze,
Rev. Gen. 3: 194. 1893; Lam, Bull. Jard. Bot. Buitenzorg, ser. 3, 7:
192. 1925, Rec. Trav. Bot. Ne"erl. 36: 524. 1939; Record, Trop. Woods
59: 36. 1939; Baehni, Candollea 9: 147. 1942; Lam, Blumea 5: 336.
1943; Cronquist, Lloydia 9: 257. 1946, Brittonia 7: 1. 1948; van
Royen, Blumea 8: 207, 235. 1957; Heermann-Erlee & van Royen,
Blumea 8: 452. 1957; van Royen, Nova Guinea N. S. 10: 134. 1959;
Meeuse, Bothalia 7: 332. 1960; Wood & Channell, Journ. Arn. Arb.
41: 11. 1960; Aubre"ville, Adansonia 1: 155. 1961; Baehni, Boissiera
11:48. 1965.

Trees or shrubs; leaves without stipules, venation variable; sec-
ondary nerves numerous and parallel or fewer and distant; flowers
in umbels or on short axillary axes; sepals (4)-5-(6), without appen-
dix; corolla thin mostly tubular, with (4) -5 -(6) lobes; staminodes
present. Ovary with generally 4-5 loculi; fruits indehiscent, 3-5-
seeded; seeds with a narrow (rarely rather broad) scar; shorter than
the seed, embryo vertical.

Calyx 4-5 merous, imbricate or quincuncial.

Leaves with very numerous lateral veins pressed together.
Blades glabrous below.

Flowers sessile (pentamerous) Pouteria cylindrocarpa.

Flowers pedicelled.

Blades less than 10 cm. long.

Flowers 4-merous Pouteria venulosa.

Flowers 5-merous Mastichodendron Williamsii.

Blades more than 12 cm. long, flowers 5-merous.

Pouteria Ulei.

152 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

Blades silky or pilose beneath, flowers pentamerous.

Blades emarginate or rounded at the end, pedicels more than

10 mm. long Pouteria peruviana.

Blades long-acuminate, flowers almost sessile.

Pouteria sanctae-rosae.
Blades obovate, short acuminate, pedicels 2-4 mm. long.

Pouteria Duckeana.
Leaves with 10-20 pairs of lateral veins, fairly separated.

Blades glabrous below, except in some instances the hairy mid-
rib and principal veins becoming glabrate with age.

Flowers sessile (pentamerous) Pouteria validinervis.

Flowers pedicelled.

Flowers pentamerous (sometimes hexamerous : Pouteria ani-
bifolia) .

Flowers — excluding pedicels — more than 15 mm. high.

Pouteria lucuma.

Flowers — excluding pedicels — decidedly lesser than
10 mm. high.

Pedicels less than 6 mm. long.

Calyx membranous, less than 2 mm. high, corolla
2.5 mm. high, with rounded lobes.

Pouteria anibifolia.

Calyx fleshy, 4 mm. high, corolla 5-6 mm. high, with
triangular lobes Pouteria Bonneriana.

Pedicels more than 10 mm. long, fascicled.

Blades with distinct network above; calyx almost
glabrous Pouteria peruviensis.

Blades with obsolete network above; calyx puberu-
lent Pouteria Aubrevillei.

Flowers tetramerous.

Pedicels very short, 1-3 mm.

Calyx glabrous or with very few hairs.

Corolla 4 mm. high.

Richardella glomerata var. glabrescens.

Corolla up to 7 mm. high Pouteria caimito.

Calyx tomentose.

Corolla 7-9 mm. high Pouteria torta.

FLORA OF PERU 153

Corolla 18 mm. high Pouteria Wurdackii.

Pedicels 5 mm. long or longer.

Pedicels and calyx glabrous, corolla 8 mm. long.

Pouteria pisquiensis,

Pedicels and calyx densely coppery pilose, corolla 2 mm.
high Pouteria loretensis.

Blades fairly uniformly hairy or silky below.

Flowers sessile, or with short and stout pedicels, 2-4 mm. long.

Flowers pentamerous, very small, 2 mm. high; network not
very distinct, flat petioles 5-7 mm. long.

Pouteria tarapotensis.

Flowers tetramerous, 5-6 mm. high; network quite obsolete,
petioles less than 10 mm. long.

Richardella glomerata var. glomerata.

Flowers pedicelled, with pedicels more than 6 mm. long; net-
work quite obsolete; petioles at least 15 mm. long.

Flowers tetramerous Pouteria cinnamomea.

Flowers pentamerous (sometimes hexamerous).

Richardella macrophylla.

Calyx pentamerous, valvate; filaments completely free.

Pouteria Bouffardiana.

Pouteria anibifolia (A. C. Smith) Baehni, Candollea 9: 269.
1942. Lucuma anibaefolia A. C. Smith, Bull. Torr. Bot. Club 60:
389. 1933. Lucuma Batten-Poollii Benoist, Bull. Soc. Bot. France
84: 636. 1938. Franchetella anibifolia (A. C. Smith) Aubr. Adan-
sonia 1: 183. 1961.

Small or medium-sized tree, up to 12 m. high, with slender, terete,
glabrous, reddish-brown branchlets, however, furrowed and opaque
just at the extremity. Leaves papery, elliptical, acuminate (in Pe-
ruvian specimens, acumen well marked, up to 10-15 mm. high), more
rarely slightly obovate, often sinuate, glabrous, dull, 12-18 cm. long,
2.5-6 cm. wide; petioles slender, canaliculate, 6-12 mm. long; mid-
rib slightly prominent beneath; secondary nerves in 14-16 pairs, very
thin, patent and arching near the margin ; tertiary nerves forming a
dense but inconspicuous network. Flowers white, fascicled by 5-10,
in the leaf axils, or more commonly on short lateral shoots, with
acute-triangular and very small bracts at the base of the slender and
puberulous, 3-5 mm. long pedicels; calyx 5-6-merous, sepals almost

154 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

free, ovate-triangular, the exterior bigger and thicker than the inte-
rior, 1-1.5 mm. high, with membranaceous and ciliate margins; co-
rolla campanulate, 2-2.5 mm. high, fleshy, with 5-6 lobes as long as
the tube, rounded and ciliate; staminodes fleshy, ovate-acute, near
1 mm. high; stamens apparently fertile, with very short filaments
inserted on the upper half of the tube, anthers acute, 0.5 mm. high;
pistil much shorter than the corolla, scarcely 1mm. high, with globose-
depressed ovary, more or less densely sericeous, 1-celled, with one
ovule attached at the apex of the loculus; style very short (less than
0.25 mm.), conical and obtuse. Fruit olive-like, 2-2.5 cm. long, 8-
15 mm. wide; seed ellipsoid, 2 cm. long, with a long and narrow scar.

Type: Krukoff 1W (Matto Grosso, Brazil).

Brazil (also cultivated), Peru. Loreto, Florida, Rio Putumayo,
at mouth of Rio Zubineta, alt. 180 m., forest, King 2363 (F, G, S)—
San Martin: Juan Jui, Alto Rio Huallaga, alt. 400 m., forest, Klug
3899 (F, S).

Pouteria Aubrevillei Bernardi, Candollea 22: 231. 1967. Ere-
moluma Wurdackii Aubr. Adansonia 5: 197, t. 1. 1965, non Pou-
teria Wurdackii Aubr., 1965.

Tall tree, up to 30 m. high, with copious latex and strong greyish
branchlets furrowed at the extremities and covered with a rusty,
short, appressed pilosity. Leaves obovate, coriaceous, subopposite
at the extremity of the branchlets, alternate elsewhere, glabrous ex-
cept for some scanty and weak hairs along the midrib, short but
markedly acuminate, attenuate toward the base and merging finally
in the flat petioles 5-10 (15) mm. long; blades 9-12 (16) cm. long,
3-5 (8) cm. wide; midrib strong-prominent beneath; 8-13 pairs of
lateral veins, slender and prominulous beneath, ascending and arch-
ing, free up to the margin; tertiary veins forming a very fine, promin-
ulous and remarkable network on both faces. Flowers above the leaf
scars in several tri-florous fascicles crowded together; pedicels slen-
der, 10-13 mm. long; calyx quincuncial, with 5 sepals connate near
the base, the two interior rounded, the others ovate-acute, pilose on
the back, puberulous inside, ciliate; corolla campanulate, approxi-
mately 5 mm. high, with 5 lobes imbricate, rounded, membranous,
2.5-3 mm. high, ciliolate, rusty puberulous on the back, becoming
glabrate, tube short, fleshy, 1.5-2.5 mm. long; 5 fertile stamens with
sessile anthers in the type (subsessile in Wurdack 2478) inserted on
the middle of the tube, 1.5-2 mm. high, apiculate; staminodes 5,
fleshy, subulate and very acute, in some instances in the same flower,

FLORA OF PERU 155

one or more expanded and petaloid, 2-3 mm. long; pistil conical,
glabrous, scarcely 2 mm. high, with a 1-celled ovary; ovule attached
by its superior half; style indistinct, stigma inconspicuous, a little
oblique, granulate. Fruit unknown.

Type: Wurdack 2333 (Dept. Loreto, Peru).

Peru. Loreto: Tree 25 m., flowers cream, rainforest on lower north
slopes of Cerro Campanquiz at Pongo de Manseriche, right bank of
Rio Marafion, alt. 300-550 m. (fl. oct.), Wurdack 2333 (F). Tree
30 m., flowers white, forested ridge on right bank of Rio Santiago
3-4 km. above mouth, alt. 300-350 m. (fl. oct.), Wurdack 2478 (LE).

Pouteria Bonneriana Bernardi, Candollea 22: 237. 1968.

Medium-sized tree, up to 15 m. high with stout branchlets, chest-
nut-brown, furrowed and shortly pilose at the extremity. Leaves
papery, obovate, less frequently elliptical, alternate to almost oppo-
site, generally with a short and broad acumen, obtuse to cuneate at
the base, glabrous above, short and loosely puberulent beneath ; mid-
rib impressed above near the petiole, prominent elsewhere, strongly
prominent beneath and, with the secondary veins, characteristically
red-brown; lateral veins in 15-20 pairs, strongly arched-ascending;
veinlets prominent on both sides, forming a dense and somewhat
irregular network, more noticeable beneath; petioles puberulent be-
coming glabrate with age, almost terete, very narrowly canaliculate
on the upper face, 35-55 mm. long; blades 20-28 (35) cm. long, 9-
11 (13) cm. wide. Fascicles 8-10 (12)-flowered in the axils of the leaf
scars, on stout and fairly woody branshlets (at least on the samples
at hand; pedicels stout, 3-4 (5) mm. long, sparsely pilose; calyx
quincuncial, with lobes almost completely free, sepals nearly orbic-
ular, fleshy, measuring 4 mm., sparsely pilose on the back, the three
outer ones hairy on the inner face of the upper margin, the inner ones
glabrous inside and with very thin margins; corolla glabrous, fleshy,
5-6 mm. high, campanulate, with lobes imbricate, half as long as the
tube, fairly triangular and auriculate at the base; stamens a little
shorter than the corolla, with easily distinguishable filaments all along
the interior of the corolla, but free only from the middle of the tube,
4 mm. long, ovate anthers 1 mm. high, basifixed; staminodes acute,
scarcely 1 mm. high; pistil 4 mm. high, silky-puberulent except
for the end of the stout style; stigma very reduced; ovary moderately
furrowed, 5-locular; the ovules attached broadly by the middle of
their adaxial side. Fruit unknown.

Type: Wurdack 24-70 (Dept. Amazonas, Peru).

156 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

Endemic to Amazonian Peru, probably present, too, in Ama-
zonian Brazil, where it could be confused with Rhamnoluma pariry
(Ducke) Baehni (=Lucuma pariry Ducke, Eglerodendron pariry
(Ducke) AubreVille) .• — Amazonas: Prov. Bagua, forested ridge on
right bank of Rio Santiago 3-4 km. above mouth, elev. 300-350 m.,
Wurdack 21*70 (G, LE). — Loreto: Prov. Alto Amazonas, rainforest on
lower southwest slopes of Cerros Campanquiz, right bank of Rio
Maranon opposite mouth of Rio Santiago, elev. 300-450 m., Wur-
dack 2514 (F).

Pouteria Bouffardiana Bernardi, Candollea 22: 228. 1968.

Tree 12 m. high known only from the type, branchlets fairly
robust, with pale grey bark, glabrous, furrowed at the ends; leaf buds
acute, white-pilose when young, finally glabrate. Papery leaves al-
most coriaceous, entirely glabrous, generally with a rounded apex,
rarely obtuse; midrib slender, slightly raised above, more conspicu-
ous and prominent beneath; lateral veins 8-12 pairs, nearly obsolete
above, slender and slightly raised beneath, spreading and gradually
arched toward the borders; network fairly inconspicuous and lax;
petioles fairly thin, shining, glabrous, half-terete (?), not at all cana-
liculate, 10-15 mm. long. Flowers 3-6 fascicled above the scars of
fallen leaves, or 1-2 flowers in the axils of young leaves at the top
of the branchlets. The flowers of the same fascicle are in different
stages of growth, from buds with short pedicels, to almost opened
flowers with elongated pedicels 2-3 mm. long and glabrous; calyx
depressed-globose, 3 mm. long and wide, peritamerous, valvate, cov-
ering completely the rest of the flower, glabrous on both sides, except
the top of the sepals very shortly and minutely white tomentose;
sepals almost equilateral triangles in shape, 1.5 mm. long each side,
in some instances irregularly opened, giving therefore the false ap-
pearance of a trimerous calyx; corolla pentamerous, 2 mm. high,
2.5 mm. wide, fleshy, glabrous with imbricate, triangular petals
slightly more than 1 mm. long; stamens 1.25 mm. long with filaments
moderately thick, nearly 1 mm. long, completely free from the tube,
inserted at the corolla base; pollen-bearing ovate anthers, with a large
connective, obtuse at the top; fleshy staminodss, barely 1 mm. long,
almost trullate; pistil depressed, scarcely 1 mm. high, 1.5 mm. wide,
ovary densely covered with silky-yellowish hairs, style robust, stigma
glandular, slightly expanded; 5-locular ovary, small and depressed
loculi, ovules attached toward the bottom of the loculi.

FLORA OF PERU 157

Loreto: Gamitana Cocha, Rio Mazan, alt. 100-125 m., Schunke
368 (Type: F, US).

Pouteria caimito (Ruiz & Pav6n) Radlk. in Sitzb. Math.-Phs.
Cl., Acad. Wiss. Munchen 12: 333. 1882, var. caimito [var. typica
Baehni, Candollea 9: 260. 1942]. Achras caimito Ruiz & Pa von, Fl.
Peruv. 3: 18, t. 240. 1802. Lucuma caimito Roem. & Schult. Syst.
4: 701. 1819; Mart. Fl. Bras. 7: 79, t. 33. 1863. Labatia caimito Mart.,
Herb. Fl. Bras. p. 170. 1837. Guapeba caimito Pierre, Not. Bot. Sa-
pot. p. 42. 1891. Lucuma ternata H.B.K. Nov. Gen. & Sp. 3: 241.
1818. Lucuma huallagae Standl. ex Williams, Field Mus. Publ. Bot.
15: 411. 1936. Pouteria leucophaea Baehni, Candollea 18: 176. 1962.

Large tree, up to 35 m., more frequently 10 m. high; branchlets
blackish, sparsely puberulous or glabrate, furrowed. Leaves variable,
in the Peruvian samples mostly chartaceous and elliptical, but also
coriaceous, obovate, acuminate or widely cuspidate, strongly atten-
uate at the base, glabrous or with a few dispersed hairs, 10-20 cm.
long, 3-6 cm. wide; petioles 5-20 mm. long, canaliculate; midrib
prominent beneath; secondary veins in 9-12 pairs, patent and arcuate,
slender; tertiary veins forming a very fine network, prominulous be-
neath. In the herbarium samples, the leaves are nearly black above,
and reddish beneath. Flowers solitary or 2-5 in the axils of leaves
or above the leaf scars, sessile or with a very short pedicel (1 mm.) ;
sepals 4 (5), ovate, glabrous inside, the outer ones smaller, up to
4 mm. long, almost glabrous on the back, the inner ones sparingly
silky, 6 mm. long; corolla cylindric, white or greenish-white, 5-8 mm.
long; lobes 4 (5), rounded, ciliate on the margin, 1-2 mm. long; stami-
nodes subulate or lanceolate, sometimes slightly ciliate, a little shorter
than the lobes; filaments attached to the middle of the tube, 2 mm.
long; anthers 1 mm. long, apiculate; pistil a little longer than the
corolla, with a small, globose ovary 2 mm. high, style 6 mm. long,
slender, stigma inconspicuous, 4-tuberculate; ovary densely rufous-
hirsute, 4-celled, with ovules attached at the bottom of the locules.
Berry edible, globose to cylindrical, obtuse or apiculate, 4-5 cm. in
diameter, tomentose when young; seeds 1-4, cylindric-ovoid, slightly
compressed on the sides, scar somewhat shorter than the seed.

Type: Ruiz & Pavdn s/n: "Habitat in Andium montibus imis
calidis et cultis ad Pozuzo et Chinchao" (Huanuco) (G).

Cultivated for the fruits in all tropical America. Loreto: Mishu-
yacu, near Iquitos, 100 m. dense forest, Killip & Smith 29993 (F).
Distrito de Iquitos, fundo Indiana, 110 m., planted, Mexia 6397

158 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

(G, S). Lower Rio Nanay, Williams 472 (F), 695 (G). Pebas on the
Amazon River, Williams 1 792 (F) . Pr6 on the Amazon River, Wil-
liams 1983 (F). Lower Rio Huallaga, Williams 4802 (F) "Lucuma
huallagae Standl." (F, G); ibidem: Williams 4717 (F, G). Santa
Rosa, lower Rio Huallaga below Yurimaguas, alt. about 135 m. dense
forest, Kittip & Smith 28778, 28798 (F).— Amazonas: Forested ridge
of Rio Santiago, 2-3 km. above mouth, 300-350 m., Wurdack 2408
(G, S). — Junin: La Merced, 650 m., Macbride 5584- Puerto Bermu-
dez, alt. about 375 m., dense forest, Killip & Smith 26564 (F) "Lu-
cuma huallagae."' — San Martin: Juan Jui, Alto Rio Huallaga, about
400 m. forest, Klug 3822 (F, S) (Pouteria leucophaea Baehni). With-
out locality, Ruiz & Pav6n (G: 3 specimens).

Pouteria cinnamomea Baehni, Candollea 9: 252. 1942. Labatia
discolor Diels, Engl. Bot. Jahrb. 37: 601. 1906, non Pouteria discolor
(Baill.) Baehni.

Tree 20 m., branchlets robust, reddish brown, a little compressed
at the ends, rusty-tomentose. Leaves papyraceous, obovate, indis-
tinctly apiculate, tapering toward the base, pubescent along the
upper face of the midrib when young, cinnamon coloured, densely
silky-tomentose below, 10-15 cm. long, 4-5 cm. wide; petioles stout,
above flat, approximately 15 mm. long. Flowers 5-6-fasciculate in
the axils of leaves, only female observed; pedicels coppery-silky,
robust, 8-10 mm. long; sepals 4, almost completely free, the two
outer thick, coriaceous, silky outside, ovate, 6x4 mm.; the inner
glabrous and only 3 mm. large, ovate-elliptical ; corolla whitish-green,
rather fleshy, 5-6 mm. long; lobes 4, ovate, margins ciliolate, shorter
than the tube; staminodes tongue-like, petaloid, fleshy in the basal
portion adnate to the corolla tube, shorter than the lobes; stamens,
in all the flowers observed, reduced to linear filaments, 3 mm. long,
attached to the inferior part of the tube; pistil pear-shaped, 5-6 mm.
high, almost completely covered by long coppery sericeous hairs,
except the top end of the style, stigma inconspicuous; ovary 1-1.5
mm. high, 4-locular, with ovules attached by the middle of their
adaxial side. Fruit not seen.

Type: Weberbauer 5030.

Known only from the type collection. Cuzco : Convenci6n, 1,700
m., Weberbauer 5030.

Pouteria cylindrocarpa (Poepp.) Baehni. Sideroxylon cylindro-
carpon Poepp. in Poepp. & Endl. Nov. Gen. & Sp. 3: 72, t. 282. 1845.

FLORA OF PERU 159

Micropholis cylindrocarpa (Poepp.) Pierre, Not. Bot. Sapot. p. 40.
1891. Pouteria saltuensis Baehni, Candollea 18: 171, fig. 60. 1962.

Shrub or small tree (up to 10 m.) ; branchlets slender, rufous-
puberulous and angled at the extremities. Leaves papyraceous, ellip-
tic, glabrous on both sides, 7-9 cm. long, 2.5-3 cm. wide, long and
abruptly acuminate (acumen up to 12 mm. long), glabrous, with very
numerous secondary, parallel veins straight and patent, spreading
from the midrib which is prominent beneath ; petioles slender, deeply
canaliculate, 8-10 mm. long. Flowers sessile or with very short ped-
icels, solitary or in pairs in the axils of leaves; sepals 5, quincuncial,
thick coriaceous, pubescent outside, puberulous inside near the edges,
2 mm. long; corolla glabrous, 2.5 mm. long, with 5 small rounded
lobes; staminodes lanceolate, half as long as the lobes, stamens small,
with very short filaments attached in the inferior half of the tube,
anthers heart-shaped; ovary 4-5-celled, hirsute, with the ovules at-
tached on the upper part of the locules. Berry cylindrical (ex Poep-
pig) or egg-shaped, apiculate, 1-1.5 cm. long, 1-celled, 1-seeded.

Dpto. Loreto: Forests of Yurimaguas, Poeppig 2371 (type G).
Mishuyacu, near Iquitos, 100 m. alt., Klug 1104 (F). Between Yuri-
magus and Balsapuerto (lower Rio Huallaga basin), 135-150 m.,
dense forest, Killip & Smith 28011 (type of Pouteria saltuensis},
28150, 28257, 29021 (F). La Victoria, on the Amazon River, Wil-
liams 2995 (F, G).

Pouteria Duckeana Baehni, Candollea 18: 161, /. 53. 1962.
Micropholis apiculata, Gilly, Sched. Herb.

Medium-sized tree, rarely attains more than 20 m. height, with
brown, stout and upright branchlets compressed, covered with a
rusty-red appressed pilosity at the extremities. Leaves fairly coria-
ceous, elliptical to obovate, more or less abruptly acuminate at the
tip, acumen from 5 to 20 mm. high, base generally acute, glabrous on
the upper side, densely red-silky beneath, becoming almost glabrate
with age; blades (12)-15-(20) cm. long, (6) 8 (12) cm. wide; petioles
stout, canaliculate, 15-25 mm. long, shortly and densely pilose; mid-
rib strong, prominent beneath; secondary veins slender, numerous,
parallel, inconspicuous. Flowers in the axils of the leaves, on very
short shoots 3-5-flowered; robust pedicels rusty-red, silky, 2-4 mm.
long; calyx quincuncial with two thick external sepals, broadly-ovate,
densely rusty pubescent on the back and pilose inside, the remainder
thinner, broadly obovate, silky, ciliate at the edges, 3-4 mm. long;
corolla tubular 4-5 mm. long, with 5 obtuse lobes, slightly auriculate,

160 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

2 mm. high; stamens attached to the throat, with short filaments;
anthers ovate, short, included; staminodes fleshy, subulate, shorter
than the lobes; pistil 3 mm. high, with depressed ovary, hirsute at
the base only, 5-celled, style stout, stigma inconspicuous; ovules at-
tached by the middle of their adaxial side. Fruit unknown.

Type: Ducke 24853, Manaos, Amazonia, Brazil (G).

Brazil, Peru. Loreto: Tierra doble on the Rio Nanay, Williams
910 (F, G).

Called Balata quebradiza in the vernacular.

Pouteria loretensis Baehni, Candollea 9: 235. 1942.

Tree (?) ; branchlets fairly robust, terete, compressed at the ends,
pubescence appressed. Leaves coriaceous, glabrous, glossy beneath,
obovate, shortly but abruptly acuminate (acumen up to 5 mm. long),
attenuated gradually toward the petiole, 10-12 cm. long, 3.5-5 cm.
wide; petioles stout, 5 mm. long; midrib fairly prominent and robust
on both sides; lateral veins very slender in 13-16 pairs, patent, only
slightly arched toward the border; veinlets and network quite obso-
lete. Flowers in fascicles of 3-5 on slender, sulcate, rusty-puberulous
branchlets 5-10 cm. long, bearing few leaves; pedicels fairly robust,
6-7 mm. long, pubescence coppery and appressed; sepals 4, 2 outer
ones thick and broadly ovate, with a coppery pilosity on both sides,
2 inner ones pubescent outside with very few sericeous hairs down the
center and with scarious and ciliate margins; corolla 2 mm. long, gla-
brous, lobes 4, rounded, slightly shorter than the tube; stamens per-
fect, with short thick filaments attached to the middle of the tube,
U-shaped, fertile anthers cordate, 1 mm. long; staminodes subulate,
subacute, half as long as the lobes; pistil almost 2 mm. high, ovary
depressed globose and coppery-silky, 2-celled, with loculi high and
narrow, ovules attached to the central portion; style fairly slender,
0.5 mm. long, stigma quite obsolete. Berry unknown.

Type : Tessmann 5^51 .

Known only from the type collection. Loreto: Middle Ucayali,
Yarina Cocha, Tessmann 5^51 (G, S).

For the vegetative organs, and also for the floral morphology,
this species is fairly similar to Pouteria lateriflora (Benth. ex Miq.)
Radlk. (= Pseudocladia lateriflora Pierre); but in Pouteria loretensis
the pedicels and flowers are notably more robust; the ovule insertion
being different also (in P. lateriflora it is completely basal), it is easy
to separate the two taxa.

FLORA OF PERU 161

Pouteria Lucuma (Ruiz & Pav6n) 0. Ktze. Rev. Gen. 3, 2: 195.
1898. Achras lucuma Ruiz & Pa von, Flora Peruv. 3: 17, t. 239. 1802.
? Lucuma bifera Mol. Saggio Chile p. 187. 1782, French ed. p. 161.
1789. ? Lucuma turbinata Mol., I.e. ? Lucuma obovata H.B.K. Nov.
Gen. 3: 241. 1818; A. DC. Prodr. 8: 172. 1844 (with var. ruizii);
Mart. Fl. Bras. 7: 69. 1863. Pouteria insignis Baehni, Candollea 9:
356. 1942. Richardella Lucuma (Ruiz & Pavon) Aubr. Adansonia
1: 175. 1961; Adansonia, Me'moire 1, p. 41, fig. 15. 1964.

Medium-sized tree, 6-15 m. high; branchlets greyish- white ap-
pressed-pilose, terete. Leaves coriaceous, rarely papery, obovate, with
generally rounded tip, in some instances subacute or obtusely acumi-
nate, acute at the base, glabrous, 10-20 cm. long, 5-8 cm. wide; peti-
oles 1.5-4 cm. long; midrib robust and prominent beneath, veins 9-
12 (14) pairs, patent or arcuate-ascending, slender. Flowers single
or by 2-3 in the axils of the leaves; pedicels stout, rusty pubescent,
8-15 mm. long; calyx pentamerous, quincuncial, with sepals ovate,
obtuse, rusty-pubescent, the inner ones thinner and with the edges
glabrate outside but ciliolate, glabrous, or slightly pubescent inside
near the tip, 8-12 mm. long, 6-8 mm. large; corolla yellowish-green,
fleshy, up to 15 mm. long, lobes 5, ovate, shorter than the tube (7-
8 mm. long), slightly puberulous outside and papillose at the margin ;
perfect stamens with filaments attached to the throat, 1 mm. free,
3 mm. adnate to the tube, anthers ovate, basifixed, 3 mm. long; stami-
nodes linear-subulate, papillose, 3 mm. long; pistil, at an thesis, as long
as the corolla, elongate-conical, ovary up to 7-8 mm. high, yellow-
sericeous at the base only (3-4 mm.), 5 (4)-locular, loculi small, locu-
lar cavities restricted to the upper portion of the ovary, ovules
attached to the top of the narrow loculi, style slender, papillose,
slightly 5-furrowed, stigma obscurely 5-tuberculate. Berry edible,
apple-shaped, apiculate or depressed, glabrous, dark green; seeds
1-5, ovoid or spherical, scar large, ovate, almost as long as the seed.

Type: Ruiz & Pavon s/n, Peru (G).

Peru, Ecuador, cultivated in Chile, Costa Rica. Loreto: along
Rio Itaya, Williams 189 (F); Tierra doble on the Rio Nanay,
Williams 901+ (F); Mishuyacu, near Iquitos, 100 m. forest, clear-
ing, Klug 1515 (F). — Cajamarca: Prov. Cutervo, Socota common
along river and in fencerows, alt. 2,800 m., Stork & Horton 10154
(F, G). — Ayacucho: Aina between Huanta and Rio Apurimac, clear-
ing, alt. 750-1,000 m., Kittip & Smith 22818 (F); Maria del Valle
probably an escape, alt. 2,300 m., Macbride 4953 (G)/ — Cuzco: 01-

162 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

lantayambo, alt. 3,000 m., Cook & Gilbert 471, 472, 474 (F).— With-
out locality: Dombey 1+10 (G); Pavon 585 and without number (G).

Pouteria peruviana Baehni, Candollea 9: 214. 1942. Lucuma
macrophylla Krause, Verhandl. Bot. Ver. Brandenburg 1908 (L): 94.
1909. Non Pouteria macrophylla (Lam.) Eyma (1936).

Tree 6 m. (collected only once) ; branchlets robust, glabrous, te-
rete, brown. Leaves thick coriaceous, oblong or obovate-oblong,
rounded or emarginate at the apex, gradually attenuated in the stout,
dilated petiole, 20-25 mm. long, subcanaliculate ; blade 16-20 cm.
long, 6-8.5 cm. wide, glabrous and shining above, coppery dense ap-
pressed pubescent beneath, becoming glabrate with age; midrib very
prominent beneath ; secondary nerves numerous, patent, parallel, slen-
der. Flowers conspicuous, 4-8 above the leaf scars, pedicels stout, 12-
16 mm. long, short rusty-pilose; sepals 5, almost completely free,
very thick-coriaceous, quincuncial, rusty-pilose on the back, glabrous
inside, 5-7 mm. long, 3-4 mm. large, thin-edged, especially the two
inner ones, corolla tubular, fleshy, 8-10 mm. long, with 5 lobes ovate,
3 mm. high; fertile stamens erect and exserted, attached to the throat,
with filaments 5 mm. long, anthers narrow, 2 mm. high; staminodes
subulate, 2 mm. long; pistil shorter than the corolla, 4-5 mm. high,
ovary conical, brown, hirsute, 5-celled, ovule attached to the top of
the loculus; style as long as the ovary, glabrous, stigma obsolete.
Fruit unknown.

Loreto: Cerro de Escalero, 1,200 m.; Ule 6793 (type: G).

Pouteria peruviensis (Aubr.) Bernardi, Candollea 22: 231. 1961.
Eremoluma peruviensis Aubr. Adansonia 5: 199. 1965.

Medium-sized tree (collected only once) up to 15 m. high with
dark branchlets sparsely puberulous at the extremity, a little com-
pressed. Leaves subcoriaceous, glabrous, elliptical and acuminate,
with acumen well marked, 10-15 mm. long, attenuate toward the
base, gradually passing into the slender, glabrous, and in part cana-
liculate petiole, 8-12 mm. long; midrib fairly strong and prominent
on both sides; secondary veins thin, somewhat irregular arched-
ascending, in 9-13 pairs, joining near the margins; frequent and
slender veinlets forming a dense, peculiar network prominulous on
both sides. Flowers fasciculate, 5-10 in the axils of the leaves, ped-
icels thin, however, thickening below the calyx, sparsely puberulous;
sepals 5, quincuncial, nearly orbiculate, measuring 3 mm., very fleshy
and thick at their base, slightly membranaceous at the borders,

FLORA OF PERU 163

sparsely pilose on the back, glabrous inside, obscurely ciliolate, im-
perceptibly connate at their bases; corolla nearly 5 mm. high, gla-
brous, membranaceous, campanulate, with 5 lobes up to 4 mm.,
nearly orbiculate, presenting dark and numerous veinlets; fertile sta-
mens 5, inserted on the throat, anthers sessile, ovoid, erect, apiculate,

1 mm. high; staminodes 5, subulate, acute, 2-3 mm. long; pistil
conical, much shorter than the corolla, 2 mm. high, glabrous, the
broad style merging with the top of the ovary, stigma punctiforme;
ovary 1-celled, with one ovule broadly attached by the adaxial face
at the upper part of the loculus. Fruit unknown.

Type: Wurdack 2368.

Known only from the type: Loreto: Alto Amazonas, rainforest
on lower north slopes of Cerros Campanquiz at Pongo de Man-
seriche, right bank of Rio Maranon, alt. 300-350 m., Wurdack 2363
(F).

Pouteria pisquiensis Baehni, Candollea 9: 263. 1942.

Medium-sized tree, with a trunk 30 cm. in diameter (collected
only once), branchlets terete, glabrous, black at the ends only, other-
wise whitish. Leaves glabrous, membranaceous, obovate, indis-
tinctly acuminate or subacute or obtuse, cuneate at the base, 8-
10 cm. long, 3-4 cm. wide, petioles thin, flat on the upper side, 1-2
cm. long; midrib scarcely prominent beneath; 8-10 pairs of arcuate-
ascending, slender lateral veins, forming with the tertiary veins a fine
network, prominulous beneath, obsolete on the upper side. Flowers
2-5 in the leaf axils; pedicels 10 mm. long, thin and little compressed
at the base, swelling gradually toward the calyx; sepals 4 (4 x 4 mm.),
the two exterior large-ovate and fleshy, glabrous, the interior ones
almost orbiculate, ciliolate, scariose near the edges; corolla tubular,
6 mm. long; lobes 4, oblong-ovate, truncate, minutely papillose, 2.5
mm. high; staminodes narrowly oblong, obtuse, papillose, scarcely

2 mm. high; fertile stamens included, with filaments flat, 3 mm. long,
attached near the base of the tube, anthers dorsifixed, narrow, apic-
ulate, 1 mm. high; pistil of the same length as the corolla, ovary
very small, only 1 mm. high, hirsute, 4-locular, with ovules attached
near the base of the loculi; style glabrous, slender, smooth, stigma
obscurely capitate. Fruit unknown.

Dept. Loreto: Ucayali territory, between 10° S and its estuary,
Rio Pisqui, alt. 150 m., Tessmann 3103, type (G).

164 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

Pouteria sanctae-rosae Baehni, Candollea 9: 211. 1942.

Tree 12-15 m., branchlets robust and terete, compressed, grooved
and covered with a dense, ochre-yellow pilosity at the ends. Leaves
oblong, coriaceous, abruptly mucronate (mucro narrow and fragile,
5-8 mm. long), cuneate at the base, glabrous and shining above,
dense, ochre-yellow pilosity below, midrib a little impressed above,
stout and very prominent below, secondary veins very numerous,
patent and straight except near the border, where they are arched,
more conspicuous beneath. Short shoots bearing many flowers, above
the leaf -scars; flowers covered with ochre-colored hairs; pedicels
less than 1 mm. long or absent; calyx pentamerous, quincuncial,
sepals almost free, 2x2 mm., the exterior coriaceous and uniformly
thick, ovate and acute, hirsute on the back, the interior ones almost
orbiculate, thinner near the ciliolate edges, glabrous inside; corolla
glabrous, fleshy, a little exserted, 3.5-4 mm. high, tubular, lobes 5,
broad-ovate, only 1 mm. high; staminodes petaloid, ovate, roundish,
almost auriculate, half as long as the lobes, filaments short, attached
to the throat, anthers not seen; pistil as long as the corolla, ovary
globose and sericeous, 2 mm. high, 5-celled, the ovules attached at
the middle or a little above. Young berry cylindrical, densely hirsute.

Type: Kittip & Smith 28799.

Known only from Peru. Loreto : Santa Rosa, lower Rio Huallaga,
below Yurimaguas, Kittip & Smith 28799 (F, G), 28094 (F).

Pouteria tarapotensis (Eichl. ex Pierre) Baehni, Candollea 9:
273. 1942. Lucuma tarapotensis Eichl. ex Pierre, Not. Bot. Sapot. 24.
1890. Franchetella sp. Pierre, I.e. Franchetella tarapotensis (Eichl.
ex Pierre) Baillon, Hist, des PI. XI: 291. 1892; AubreVille, Adansonia
1: 183. 1961.

Tree or small shrub with slender, red-brown branchlets compressed
and appressed- — with sericeous extremities. Leaves chartaceous, obo-
vate-acuminate, attenuate toward the cuneate base, undulate at the
margin, sparingly pilose or glabrate above, more or less densely silky
or pilose below, 9-13 (16) cm. long, 3.5-5 (7) cm. wide; petioles semi-
terete, silky, 5-7 mm. long; midrib moderately prominent below, 10-
12 pairs of secondary veins ascending and arching toward the margin,
prominulous beneath; network of tertiary veins more or less loose and
prominulous on both sides. Flowers crowded 5-8 in the axils of the
leaves, quite small, rusty pilose, subsessile or with indistinct and very
short pedicels, less than 0.5 mm. long; calyx quincuncial with sepals

FLORA OF PERU 165

connate for one-third of the entire length, appressed rusty-silky on
the back, ovate, acute, thick, the two outermost pilose inside near the
extremity, the others glabrous inside, nearly 2 mm. high; corolla
tubular, thin, glabrous, only 2 mm. high, with 5 rounded lobes as
long as the tube; fertile stamens (in the type) with short filaments
inserted halfway up the tube, or a little higher, anthers ellipsoid or
ovoid, 0.5 mm. high, staminodes triangular, fleshy, scarcely 0.5 mm.
high; pistil only 1 mm. high, glabrous, with base enlarged, fleshy
and ring-shaped when young, nearly conical later, with one-celled
ovary and one ovule inserted at the apex of the loculus; style atten-
uate to the small granulose stigma. Berry oliviform, 20-25 mm.
high, 7-9 mm. wide, brown appressed pilose, with navicular seed com-
pressed, 18 mm. high, 7 mm. broad with scar completely covering the
compressed adaxial face.

Type: Spruce 4561.

Peru (Ecuador, Brazil ?). Loreto: Between Yurimaguas and
Balsapuerto, lower Rio Huallaga basin, alt. 135-150 m., dense for-
est, Killip & Smith 28228 (F) . Lower Rio Huallaga, edge of forest,
Williams 3876 (F). Near Tarapoto, Spruce 4561 (G, F: fragment).

Pouteria torta (Mart.) Radlk. Sitzb. Math.-Phys. Cl. Akad.
Wiss. Munchen 12: 333. 1882. Labatia torta Mart. Herb. Fl. Bras.
174. 1837. Lucuma torta (Mart.) A. DC. Prodr. 8: 167. 1844; Mart.
Fl. Bras. 7: 74, t. 31. 1863. Guapeba torta (Mart.) Pierre, Not. Bot.
Sap. 43. 1891. Lucuma dolicophylla Standley ex R. E. Shultes, Bot.
Mus. Leafl. Harv. Univ. 13: 289. 1949; Field Mus. Publ. Bot. 15:
411. 1936.

Ligneous plant very variable in size from a little shrub to a tree
up to 25 m. high ; branchlets robust, greyish or rusty tomentose, be-
coming glabrate, with large and prominent leaf scars, terminal bud
woolly. Leaves elliptical or obovate-oblong, papery, with a very
variable extremity: rounded, obtuse, or acute, the base slightly
rounded or subacute, sparsely puberulous above, becoming glabrate
and shining, frequently greyish or rusty-pubescent beneath, more
densely along the veins, the tomentum not persistent; in several in-
stances, adult leaves were almost glabrous; midrib very strong and
prominent beneath, lateral veins in 14-28 pairs, patent-arched, pro-
minulous above, prominent beneath; tertiary veins forming a dense
and somewhat regular network; blades 12-15 (30) cm. long, 6-12 cm.
wide; petioles strong, ligneous, terete or semiterete, never canalicu-
late, 10-30 mm. long. Flowers in the axils or above the leaf scars,

166 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

frequently almost completely covering the end of the branchlets,
rusty-tomentose all over; pedicels short, 1-2 mm. long; calyx tetram-
erous, sepals almost free, the two exterior ovate, 4-5 mm. high,
tomentose on the back, glabrous inside; the interior, large elliptical,
up to 6 mm. long, sericeous on the back, glabrous inside, ciliolate,
glabrate and thinner near the borders; corolla tubular, glabrous, 7-
9 mm. long with lobes rounded, 2-3 mm. high, patent, ciliolate;
staminodes triangular acute or subulate, half as long as the lobes;
stamens included in the corolla, with filaments attached to the mid-
dle of the tube, 2-3 mm. long, with anthers 2-5 mm. high; pistil a
little longer than the corolla, with ovary small, globular, 1-2 mm.
high, hirsute with long and erect hairs, (3)-4-celled, ovules attached
to the bottom of the loculi ; style slender, 8 mm. long, stigma obsolete.
Berry ovoid, obtuse, somewhat wrinkled, short-hirsute, 3-3.5 cm.
long, 2-2.5 cm. wide, one seeded, nearly 2 cm. long, with scar 5 mm.
wide and as long as the seed.

Type: Martins s/n, Serro Frio, Minas Gerais, Brazil.

Brazil, Peru, Paraguay, Bolivia. Loreto: Timbuchi on the Rio
Nanay, Williams 886 (F). Lower Rio Huallaga, Williams 4951 (F).
Region of the Rio Ucayali from 10° South to its mouth, Tessmann
3192 (G, S). Gamitana Cocha, Rio Mazan, alt. 100-125 m., J. M.
Schunke 56 (F). Rio Jurua (Peru or Brazil ?), Ule 5163 (G).— Hua-
nuco: Prov. Huanuco, Las Palmas, river shore, Asplund 12968 (S).

Pouteria Ulei (Krause) Baehni, Candollea 9: 217. 1942. Side-
roxylon Ulei Krause, Verhandl. Bot. Ver. Brandenb. 50: 95. 1909.
Micropholis Ulei (Krause) Eyma, Rec. Trav. Bot. Ne"erl. 33: 198.
1936. Sideroxylon Quinilla Standl. ex Williams, Field Mus. Publ.
Bot. 15: 414. 1936. Pouteria Klugii Baehni, Candollea 14: 76. 1952.

Tree, tall or medium-sized; branchlets slender, terete, glabrous,
greyish; leaves membranous, glabrous, elliptical or oblong-elliptic,
in some instances obovate, sharply acuminate, with a 10 mm. long
point, blades 8-15 cm. long, 4-5 cm. wide, frequently undulate at the
edges, petioles 8-12 mm. long, deeply canaliculate; midrib strong
prominent beneath, lateral veins very numerous, parallel and slender,
filling the entire surface. Flowers in few-flowered fascicles (2-5) in
the axils of the leaves or leaf scars; pedicels slender, 4-6 mm. long,
covered with a rusty-red pilosity; calyx quincuncial, sepals 2.5 mm.
high and wide, with very thick outer ones, broadly ovate, with rusty-
red pilosity on the back, glabrous inside; inner sepals very thin, gla-
brous along the edges; corolla glabrous, tubular-campanulate, almost

FLORA OF PERU 167

3 mm. high, with 5 rounded lobes, shorter than the tube; 5 stamens
inserted on the throat, with very short filaments widened at the base,
and small anthers; staminodes fleshy, ovate, shorter than the lobes;
pistil conspicuous, conical, almost completely glabrous, 3-3.5 mm.
high, with ovary quite indistinct from the large style, which ends in
the small, obscurely 5-lobed stigma; the ovary, however, is densely
hairy, 5-celled; loculi small and narrow, ovules attached by the
upper part of their adaxial side (face). Fruit, in the herbarium
samples, almost woody, ovoid, light brown and opaque, 25 mm. high,
18 mm. wide, with 1-2 (more ?) seeds 20-23 mm. long, 6-8 mm. wide
and 3 mm. thick; scar narrow (3 mm.) and extended for the entire
length of the seed.

Type: Ule 51 6 a: Brazil, Rio Jurua, Marary (G).

Brazil, Peru. Loreto: Vicinity of Iquitos, hacienda Soledad on
Rio Itaya, inundation belt of the river, Asplund 14-4.85 (S). Mishu-
yacu, near Iquitos, alt. 100 m., Klug 130 (F, G). Along Rio Itaya,
Williams 20, 191 (F). Santa Rosa, lower Rio Huallaga below Yuri-
maguas, dense forest, alt. 135 m., Killip & Smith 28801 (F). Lower
Rio Huallaga, Santa Rosa, Williams 4903 (F, S), idem. Williams
5089a (F).

Pouteria validinervis (Sleumer) Baehni, Candollea 9: 363. 1942.
Lucuma validinervis Sleumer, Notizbl. Berlin 15: 383. 1941.

Medium-sized to tall tree (ex Klug) up to 25 m. (ex typo) ; branch-
lets robust, angular-compressed and puberulous at the extremities.
Leaves obovate, shortly acuminate or rounded or even slightly emar-
ginate, cuneate at the base, subcoriaceous, entirely glabrous, with
blades 12-18 cm. long, 5.5-8 cm. wide, petioles glabrous, stout, quite
flat above, 2-2.5 cm. long ; midrib large and prominent beneath, lat-
eral veins in 11-14 pairs, straight and almost parallel, except near
the border where they are curved ; tertiary veins starting from the
midrib at almost 90°, uninterrupted, forming a very slender but pe-
culiar reticulation. Flowers cream-colored, in the axils of leaves or
above the scars, clustered by 5-10, almost sessile, or with a very short
pedicel; calyx pentamerous and quincuncial, with very thick external
sepals, almost orbiculate and rusty-pilose on the back, glabrous in-
side, 3 mm. wide; the interior ones, smaller (2 mm. wide), thinner
and with scarious border; corolla fleshy, tubular-campanulate, near
to 4 mm. long, with 5 lobes quincuncial, ovate and obtuse, glabrous
at the margin, a little longer than the tube; staminodes subulate,

168 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

almost as long as the lobes; stamens fertile with filaments 1.5 mm.
long, enlarged at the base, attached to the throat, anthers oblong,
apiculate, 1 mm. long, extrorse, dorsifixed, included; pistil a little
shorter than the corolla, 3 mm. high, ovary ovoid, puberulous, 5-
celled, ovules attached adaxially at the middle; style stout, glabrous,
furrowed, stigma slightly enlarged and warty. Fruit unknown.

Ecuador, Peru. San Martin: Juan Jui, Alto Rio Huallaga, alt.
400 m., forest, King 3785 (F, S).

The Peruvian plant differs in several points from the description
of the type (Herta Schultze Rhonhoff 2873, Ecuador), cf. Sleumer I.e.
In the opinion of the late Prof. Baehni, however, these small differ-
ences do not justify a varietal segregation.

Pouteria venulosa (Mart. & Eichl.) Baehni, Candollea 9: 195.
1942. Sideroxylum venulosum Mart, et Eichl. ex Miq. in Mart. Fl.
Bras. 7: 52, t. 20, fig. 2, t. 37, fig. 4- 1863. Micropholis venulosa (Mart.
& Eichl.) Pierre, Not. Bot. Sap. 40. 1891. Micropholis calophylloides
Pierre, I.e.; Dubard, Ann. Mus. Col. Marseille 20: 71. 1912. Meio-
luma guyanensis Baill. Hist. PI. 11: 284. 1891. Micropholis mucro-
nata Pierre in Urb. Symb. Ant. 5: 112. 1904. Pouteria polyneura
Baehni, Candollea 7: 133. 1936; ibid. 9: 194. 1942. Pouteria flava
Baehni, Candollea 18: 164, fig. 56. 1962. Xantolis venulosa (Mart. &
Eichl.) Baehni, Boissiera 11: 24. 1965.

Medium-sized or tall tree, from 15 to 40 m. high (60 m.? see Kru-
koff 6657) ; branchlets slender, rough, greyish, compressed and rusty
pilose at the ends. Leaves chartaceous, elliptic to oblong-elliptic, some-
times ovate, rarely obovate, long acuminate, base variable, glabrous
except on the main nerve rusty pilose below, 4-10 (12) cm. long,
2-4 cm. wide, petioles rusty pubescent, slightly canaliculate, 2-5 mm. ;
midrib very thin, raised above, and fairly prominent below, lateral
veins thin, numerous, parallel, close together, a continuous marginal
nerve is present along the whole border. Flowers fascicled 6-10 (20)
in the leaf axils, with brown-silky pedicels 2-5 mm. long; calyx
tetramerous with almost completely free, large ovate, almost orbic-
ular sepals, the external thicker and bigger, 2 mm. high, hairy on the
back and ciliolate, glabrous inside; corolla tetrameous, fleshy and
glabrous, 2-3 mm. long, lobes rounded, slightly shorter than the tube ;
filaments very short, attached to the throat, anthers cordate, very
small; staminodes ovate, half as high as the lobes; pistil shorter than
the perianth, ovary globular, hirsute, style approximately the same

FLORA OF PERU 169

length, stigma obsolete; ovary 4-celled, ovules attached at the middle
of their ventral side. Berry ellipsoid, mucronulate, woody, 10-20 mm.
long, 8-10 mm. thick; seed one, ovoid, laterally compressed; scar
linear, as long as the seed.

Type: Spruce 3506 "Ad flumen Guainia vel Rio Negro supra
ostium fluminis Casiquiari," Venezuela.

Colombia, Venezuela, Guianas, Brazil, Peru. Loreto: Mishuy-
acu, near Iquitos, 100 m. alt., forest, Klug 1376 (F). Palta Cocha on
the upper Rio Nanay, Williams 3187 (F).- — Huanuco: Prov. Pachi-
tea, Distr. Honoria, Salazar 1 (F) .

I agree with the arguments of Prof. AubreVille about the genus
Xantolis (cf. Adansonia 6: 191-192. 1966) and I think that this spe-
cies, on the basis of its floral morphology, cannot be included in the
old world genus Xantolis (see also: van Royen, Blumea 8: 207-233.
1957 and AubreVille, Flore du Cambodge etc. n° 3: 74-83, 1. 13. 1963).

However, the controversial genus Micropholis, as envisaged re-
cently by the same Prof. Aubreville (Adansonia 1: 176. 1961) (and
also by Cronquist, Lloydia 9: 252. 1946) would comprise only species
with pentamerous flowers; Pouteria venulosa on the other hand is
strictly tetrameous — contrary to the inexact statements of Miquel
in Martius, Fl. Bras.; thus, the species in question, according to
Aubre"ville's arguments, is not Micropholis, though all the vegetative
characters point toward that concept. The genus Paramicropholis
Aubr. et Pellegr. (Adansonia 1: 171. 1961) has tetrameous flowers,
but quite different fruits; the old genus Gomphiluma Baill., placed
by the late Prof. Baehni in Pouteria, has also tetrameous flowers and
leaves with very numerous parallel secondary veins. The generic
concepts, founded on the vegetative characters (as Micropholis) or
in the tetramerous or pentamerous flowers, seems to me to be a hope-
less cause, especially in this family of endless modifications in the
status of the species.

If Micropholis should be conserved, with flowers tetra- and pen-
tamerous, then Pouteria must also be on the same basis and not re-
served for tetramerous species as Prof. Aubreville would have it.
Finally, Micropholis would only rest on the peculiarity of the lateral
veins; a character which appears here and there throughout the whole
family Sapotaceae, and is considered to be of sectional and not of
generic value. For all these reasons, I think that this species should
better be considered in the genus Pouteria, sensu Baehni.

170 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

Pouteria Wurdackii (Aubr.) Adansonia 5: 203, t. 2. 1965.

Tree 15 m. high (collected only once) with very stout branchlets
(13 mm. diameter in botanical samples) rusty pubescent at the ends,
with tomentose buds. Leaves coriaceous, elliptical, shortly acumi-
nate (acumen 5-7 mm. long), cuneate at the base; blades up to 30 cm.
long, 12-15 cm. wide, above almost completely glabrous, beneath
rusty pubescent along the principal veins, puberulous — becoming
glabrate — elsewhere; midrib slender above, very strong and promi-
nent beneath; 25-30 pairs of lateral veins prominent and conspicuous
beneath, arching toward the margin: tertiary veins prominulous be-
neath, forming a dense and irregular network; petioles stout, terete,
pubescent when young but later glabrate, 2.5-3 (5) cm. long. Flowers
sessile above the leaf scars, very numerous all along the terminal
branchlets, with concave and ciliolate bracteoles, 2 mm. high at the
base; calyx tetramerous, with sepals quite free, the two exterior con-
cave-ovate, rusty-pilose on the back, glabrous inside, thick, 7-9 mm.
long, 6-7 mm. wide; the interior up to 12-13 mm. long, with the mar-
gins thinner and glabrous, ciliolate at the borders; corolla glabrous
and white, fleshy, tubular, up to 18 mm. high, with 4 lobes rounded
and minutely ciliolate, 5 mm. high; stamens 4, filaments semi-terete,
inserted on the middle of the tube or a little above, anthers basifixed,
cordate, slightly apiculate, 2-2.5 mm. high; staminodes 4, subulate,
near to 5 mm. long; pistil as long as the corolla, ovary densely hir-
sute, hairs straight, beneath the tomentum deeply sulcate and trun-
cate, only 2 mm. high, 4-celled, ovules attached near the bottom of
the loculi. Berry unknown.

Loreto: High rainforest along Rio Maranon near Teniente Pinglo,
just above Pongo de Manseriche, alt. 250-300 m., Wurdick 2115
(Typus: F). Tierr£ Doble on the Rio Nanay, Williams 899 (F).

PRIEURELLA Pierre

REFERENCES: Baillon, Histoire de Plantes 11: 297. 1891-1892;
Engler in Engl. et Prantl, Nat. Pflanzenfam. Nachtr. p. 278. 1897;
AubreVille, Adansonia 1, 19. 36. 1961, id. 4, 369-371. 1964; Baehni,
Boissiera 11: 77. 1965.

Small American genus of four species of small or medium-sized
trees encountered, up to date, in the Amazonian region (sensu lato) ;
coriaceous leaves, generally oblanceolate, glabrous or more frequently
with a rusty pilosity. Inflorescences cauliflorous or on the aged and
fairly wooded branches; flowers (4) 5-merous, with slender and rela-

FLORA OF PERU 171

tively long pedicels; calyx quincuncial, sepals almost completely free;
corolla broadly campanulate, without appendages, tube very short;
stamens attached to the throat, staminodes none; pistil with obso-
lete style, ovary (4) 5 locular, berry 4 (5) seeded, seed with short and
narrow scar.

This small genus is based on a single character that is not very
impressive. I think it is doubtful if the genus can really be separated
from Chrysophyllum in the broad sense by either fruits or seeds. The
cauliflorous inflorescences, in comparison to those borne on the
branches in Chrysophylllum, might be used to differentiate Prieurella.

However, in the Bignoniaceae, Lecythidaceae, Meliaceae, Ster-
culiaceae and in the genus Swartzia, for instance, the cauliflorous
species are firmly maintained in the same genus to which ramiflorous
species pertain.

Prieurella Prieurii (A.DC.) AubreVille, Adansonia 4: 370. 1964.
Chrysophyllum Prieurei A.DC. Prodr. 8: 161. 1844. Ecclinusa Pri-
eurii (A.DC.) Aubre>ille, Adansonia 1: 20. 1961.

Tree medium-sized, with hard wood; branchlets robust, com-
pressed, furrowed and with rusty-red, appressed pilosity at the ends.
Leaves very coriaceous, obovate, rounded, emarginate or broadly
acute at the extremities, sharply cuneate at the base; petioles robust,
terete, 3-4.5 cm. long; blades 13-18 cm. long, 8-10 (11) cm. wide,
glabrate and shining above, with a dense, shortly appressed rusty-red
pubescence beneath; midrib very thin but prominent above, strong
and prominent beneath; lateral veins almost completely straight,
arching slightly only near the edges, in 10-12 pairs, fairly promi-
nent beneath, quite flat above; secondary veins nearly orthogonal to
the midrib, generally uninterrupted from one to the next; veinlets
scarcely noticeable beneath, hidden by the hairs. Flowers in fascicles
of 5-10 in the axils of the leaf scars on fairly mature (2-3-year-old)
branchlets, 15 mm. diameter, and also — according to the collectors—
on the old branches and trunk; pedicels slender, 10-15 mm. long, fur-
rowed and compressed, slightly enlarged toward the top, pilosity
dense, composed of minute rusty-red hairs; flower-bud globose, 3 mm.
wide; calyx quincuncial, with sepals almost completely free, keeled,
densely silky inside, pilose on the back, the outer two coriaceous,
large ovate, 3x3 mm., with entire margins, the inner two elliptical,
3x2 mm., long ciliolate; the intermediate sepal somewhat irregular
in shape, acute, the covered portion long, ciliolate; corolla pentam-
erous, imbricate, broadly campanulate; petals glabrous, obscurely

172 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

keeled, broadly elliptical, 3 x 2.5 mm., connivent at the callose base
for 1 mm.; stamens short, 2 mm. high, with flat filaments attached
to the corolla throat, anthers ovate, dorsifixed, extrorse, apiculate;
staminodes none; pistil silky pilose, deeply furrowed, style obsolete,
stigma obscurely 5-lobed; ovary 5-celled, ovules attached to the top
of the loculi by the upper part of their adaxial side. Fruit not extant.

Guianas, Amazonian Brazil and Peru. Huanuco: Distr. Alomia
Robles, Maroma Alta, Hacienda Delicias, alt. 800 m., Gutierrez
114 (F).

The Peruvian sample exhibits bigger leaves and petals that are
completely glabrous inside, while in the type of this species (Leprieur
s/n, Hb. DC, G) a tuft of red-brown hairs is noticeable at the base
of the inner face of the petals. However, I think that these differ-
ences are not sufficient to warrant a distinct specific or varietal status.

Prieurella Wurdackii AubreVille, Adansonia 5: 203, pi. 3. 1965.

Tall tree, up to 30 m. high (collected only once) with stout branch-
lets. Leaves coriaceous, obovate, shortly acuminate with a blunt acu-
men 5 mm. high, cuneate at the base, pedicels slender, semi terete,
smooth, 20-25 mm. long; blades 14-18 (20) cm. long, 5-7 (8) cm.
wide, glabrous and shining above, with silky and very scarce short
hairs beneath; midrib slender above, strong prominent beneath, lat-
eral nerves arched-ascending in 10-12 pairs; veins connecting the
nerves regularly and in an almost parallel manner; tertiary veinlets
numerous and orthogonal to the veins; the network fairly noticeable
on both sides. Many-flowered (15-20) fascicles frequently pressed
together on the fairly woody, quite defoliated branchlets of 2-3
years; pedicels very slender, slightly compressed, covered with a
sparse, short, appressed pilosity, 6-10 mm. long; flowers globose
when in the bud, 3 mm. wide; calyx quincuncial, with sepals com-
pletely free, the two outer concave, thick, almost orbicular, slightly
keeled, 2 mm. wide, sparsely pilose on the back, densely silky inside,
the two inner ovate, thinner, wide elliptical, 3x2 mm., ciliolate at
the margins, silky on both sides; the intermediate sepal has its cover-
ing half like the outer sepals and the covered half the inner; corolla
glabrous, imbricate, pentamerous, with petals almost free, nearly
orbiculate, 3 mm. high; stamens generally aborted (I did not observe
any anthers), filaments slender, 1 mm. long, inserted at the corolla
base; staminodes none; pistil short and thick, 2 mm. high, style obso-
lete, stigma whitish with 5 obscure lobes, ovary 5-furrowed, densely
silky, 5-celled, ovules attached toward the base of their adaxial side.
Fruit unknown.

FLORA OF PERU 173

Known only from the type. Loreto: Prov. Alto Amazonas, rain-
forest on lower northwest slopes of Cerro Campanquiz, Rio Maranon
just above Pongo de Manseriche, elv. 250-350 m., tree 30 m. with
copious milky latex. Flowers yellow-brown, Wurdick 2293, l type

(F, G).

RICHARDELLA Pierre

REFERENCE: AubreVille, Adansonia 1: 174. 1961; Baehni, Bois-
siera 11: 95. 1965.

Medium-sized, rarely tall trees or shrubs, flowers in fascicles, calyx
simple, generally 4-5 (6)-merous, petals thin, without appendages,
staminodes always present; ovary 4-5-celled (rarely more or less),
berry usually 1-seeded, the seed bearing a long and very wide scar,
embryo upright.

Flowers with very short pedicels; corolla lobes 4.

Adult leaves silky beneath R. glomerata var. glomerata.

Adult leaves glabrous beneath R. glomerata var. glabrescens.

Flowers with pedicels 14-18 (25) mm. long; corolla lobes 5 (6).

R. macrophylla.

Richard el la glomerata (Miq.) Baehni var. glomerata. Boissi-
era 11 : 97. 1965. Lucuma glomerata Miq. in Mart. Fl. Bras. 7 : 81, t. 36,
jig. 2. 1863. Pouteria glomerata (Miq.) Radlk., Sitzb. Math. Phys.
Cl. Acad. Wiss. Munchen 12: 333. 1882. Abatia glomerata (Miq.)
Radlk., I.e. 14: 451. 1884. Guapeba 1 glomerata (Miq.) Pierre, Not.
Bot. Sapot. 43. 1891. Labatia parinarioides Radlk., I.e. 14: 451. 1884.
Guapeba ? parinarioides (Radlk.) Pierre, I.e. Pouteria Weddeliana
Pierre, I.e. 45.

Small tree up to 12 m., or shrub 3-4 m. high, with stout, red-brown
branchlets, almost cylindrical in the lower portion of the samples,
slightly furrowed and grey puberulous at the ends. Leaves sub-
coriaceous, obovate-elliptic or obovate oblong, shortly acuminate,
generally obtuse at the base, glabrate above, silky beneath, 6-12 cm.
long, 2.5-5 cm. wide, midrib large and flat on both sides, almost
obsolete above, red or dark beneath, then showy from the whitish
surface; petioles short, 4-8 mm., stout, silky and flat above. Flowers
5-10 in axillary fascicles, tetramerous, pedicels 2 mm. long, sepals
largely almost free, elliptical, the two external thick and densely

1 In Adansonia, I.e. it is stated to be Wurdack 2290. This is a misprint.

174 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

pilose outside, glabrous inside, 4x3 mm.; the internal glabrous,
slightly narrower and ciliolate; corolla large-tubular, 4 mm. high,
with four rounded lobes only 1 mm. high, minutely ciliolate; stami-
nodes petaloid, inserted between the lobes, rounded or truncate, half
as high as the lobes; stamens 4, reaching the top of the corolla, fila-
ments inserted near the base of the tube, 2 mm. long, anthers apicu-
late, basifixed, 1 mm. high; pistil 3 mm. long, ovary long hirsute,
apparently 2 mm. high, but deprived of the hairs, only 1 mm. high,
style slender, stigma quite obsolete; ovary 4-celled, the ovules at-
tached by the middle of their ventral face. Berry apple-like, 3 cm.
in diameter; seeds 3-4, ovoid with a very wide scar.

Type: Pohl 2/n, "ad Rio Maranhao" Brazil.

Argentina, Brazil, British Guiana, Paraguay, Peru. Loreto:
Ucayali Basin, Yarina Cocha, alt. 150 m., Tessmann 3470 (G, S).

Richardella glomerata (Miq.) Baehni var. glabrescens Huber-
Pouteria glomerata (Miq.) Radlk. var. glabrescens Huber, Bull. Soc.
Bot. Geneve, ser. 2, 6: 197, fig. 11. 1914.

Tree, with elliptical membranous leaves (in Peruvian sample)
8-10 cm. long, 2.5-3.5 cm. broad with nine pairs of thin secondary
veins, glabrous when adult; flowers identical to those in the var.
glomerata. Fruit unknown.

Type: Ducke 7921, Para, Rio Cumina, near to Lake Castanho
(Brazil).

Brazil, Peru. Loreto: Region of the Rio Ucayali, from 10° S to
its mouth, Tessmann 3^86 (G, S).

Richardella macrophylla (Lam.) AubreVille, Adansonia 1 : 175.
1961. Chrysophyllum macrophyllum Lam. Tabl. Encycl. 2: 44. 1793.
Pouteria macrophylla (Lam.) Eyma, Rec. Trav. Bot. Ne"erl. 33: 164.
1938; Baehni, Candollea 9: 404. 1942. Lucuma rivicoa Gaertn. f.,
Carp, suppl. 3: 30. 1807; Miquel in Martius, Fl. Bras. 7: 71, Tav. 29.
1863. Vitellaria rivicoa Radlk. Sitzb. Math.-Phys. Cl. Acad. Wiss.
Munchen 326. 1882. Richardella rivicoa (Gaertn. f.) Pierre, Not. Bot.
Sapot. 19. 1890. Vitellaria glaucophylla Engl. Bot. Jahrb. 12: 513.
1890. Lucuma rivicoa var. glaucophylla (Engl.) Dubard, Ann. Muse"e
Col. Marseille, 2 ser. 10: 15. 1912. Lucuma acreana Krause, Notizbl.
Bot. Gart. Berlin 6: 169. 1914.

Shrub or tree up to 20-40 m., sometimes planted in tropical Amer-
ica for its edible fruits; branchlets terete and almost glabrate in the
lower portion of the samples, angulate-compressed and frequently

FLORA OF PERU 175

densely appressed rusty pilose at the ends. Leaves papery, elliptical
or obovate, frequently obtuse, sometimes acute at the extremity;
base acute, merging in the long (15-25 mm.) slender, terete, rusty-
pilose, sometimes slightly canaliculate petioles; blades glabrate
above, glaucous or whitish beneath, silky puberulous becoming fre-
quently glabrate, 14-18 (rarely more) cm. long, (4) 6-8 cm. wide;
midrib semiterete, slender, prominent beneath, 12-15 pairs of lateral
nerves, parallel, more or less patent, straight; tertiary veins almost
obsolete. Flowers in axillary fascicles of 10 or more, sometimes very
close all along the defoliate branchlets, slender pedicels 10-15 mm.
long, minute ochre-yellow pilosity; sepals 5 (6) imbricate, ochre-
yellow pilosity, the exterior generally smaller and thicker, ovate, up
to 7-8 mm. long, 5-6 mm. wide, ciliolate; corolla tubular, fleshy,
greenish-white, rather long, 8-11 mm., sparsely silky outside, becom-
ing glabrate, 5 (6) rounded lobes quincuncial, ciliolate, 4-5 mm. high;
staminodes subulate, almost 3 mm. long; stamens fertile 5 (6) with
filaments subulate almost 3 mm. long; inserted at the same height
as the staminodes, 2 mm. long; anthers basifixed, nearly ovate, 1.5-
2 mm. high; pistil as long as the caducous corolla, style projecting
in the older flower; ovary short rusty-pilose, globose, 2 mm. high,
cells 5-6, rarely more; loculi very small and opened toward the style
base; ovules attached at the top. Berry globose, 1-seeded; seed ovoid,
25-30 mm. long; scar as long as the seed, very broad.

Widely distributed in Guiana, Brazil, Ecuador, Peru, and Bolivia.

Type: Aublet s/n (P).

Madre de Dios: Seringal Auristella, Rio Acre: Ule 9691 (G).

SARCAULUS Radlk.

REFERENCE: Engler, Bot. Jahrb. 12: 508. 1890; in Engler et
Prantl, Nat. Pflanzenf. 4, 1: 142. 1891; Lam. Bull. Jard. Bot. Buiten-
zorg, ser. 3, 7: 192. 1925; Eyma, Rec. Trav. Bot. Ne"erl. 33: 192. 1936;
Cronquist, Bull. Torr. Bot. Club 73: 465. 1946; Baehni, Boissiera 11:
48. 1965.

Trees, leaves without stipules, secondary veins distant; flowers in
fascicles; calyx simple, 5-merous, imbricate or quincuncial; corolla
fleshy, very thick, globose, lobes subvalvate in appearance but really
slightly imbricate; staminodes 5, thick; fruit incompletely known,
1-seeded, scar shorter than the seed.

Two species in South America.

176 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

Blades membranous, glabrate or with scattered hairs below, network
almost obsolete, acumen 10 mm., petioles scarcely 10 mm. long;
sepals 2 mm. high Sarcaulus brasiliensis.

Blades coriaceous, puberulent below, network conspicuous, acumen
15-20 mm., petioles 15 mm. long; sepals 4 mm. high.

Sarcaulus Wurdackii.

Sarcaulus brasiliensis (A.DC.) Eyma, Rec. Trav. Bot. Ne"erl.
33: 192. 1936. Chrysophyllum macrophyllum Mart. Herb. Fl. Bras.
175. 1837, non al. Chrysophyllum brasiliense A.DC., Prodr. 8: 156.
1844. Sarcaulus macrophyllus (Mart.) Radlk. Sitzber. Bayer. Akad.
Wiss. 12: 293, 310. 1882.

Tree, branchlets robust terete and glabrous below, subangular
and fuscous above with appressed hairs. Leaves narrowly ovate to
narrowly obovate, acuminate at the tip, cuneate at the base, glabrous
and opaque above, with scattered hairs below when young, becom-
ing glabrate with age, chartaceous or coriaceous; midrib prominent
beneath, veins in 8-10 pairs arcuate, spreading, slender, obsolete
above, prominulous below, tertiary veins very slender; blades 10-
18 (22) cm. long, 5-7 cm. wide; petioles 6-10 mm. long (in the Pe-
ruvian specimens), stout, subcanaliculate. Flowers in clusters of
3-6 (8) in the axils of the leaves; pedicels filiform, broader below the
flowers, rusty-pubescent, 12-24 mm. long; sepals 5, patent, ovate,
keeled, obtuse, finely pubescent on the back, ciliolate at the margins,
glabrous inside except for a few hairs at the base, 2 mm. long and
wide; corolla globose, fleshy, very thick, appressed silky outside,
lobes 5, almost as long as the tube, scarcely imbricate, 3-4 mm. long,
stamen with filaments very short, pilose at the base, attached to the
throat, anthers emarginate, staminodes widely triangular to subulate,
on the throat, silky outside, glabrous inside; ovary 5-celled, bottle-
shaped, 5-angled, style very short, stigmas 5, punctiform and incon-
spicuous; ovules attached to the base of the loculi. Fruit not seen.

Type: In silvis ad Para et fluvium Japura, Martins (M).

French Guiana, Surinam, Brazil, Peru. Loreto: Ucayali basin,
between 10° S and its mouth, Tessmann 3241 (G, S), Tessman 34.57
(G).- — Amazonas: Rainforest along Rio Santiago, 10-15 km. above
the mouth, elev. 250 m., bushy tree 15 m. with milky latex, corolla
pink, Wurdack 2494 (F, G, LE).

Sarcaulus Wurdackii Aubr. Adansonia 5: 240. 1965.

Tree, 15 m. high (in the type) branchlets robust, cylindrical, at

FLORA OF PERU 177

the apex rufous-pubescent, with appressed, dense hairs. Leaves ellip-
tical, coriaceous, distinctly acuminate at the tip (acumen up to 15
mm. long), the base subacute or obtuse, glabrous above except for
the principal veins densely covered with reddish short and appressed
hairs, rufous-puberulous below, more densely on the veins, petioles
robust, rufous-pubescent, up to 15 mm. long, slightly canaliculate;
midrib strongly prominent below, secondary veins in 12-15 pairs,
fairly prominent below, arcuate-ascending, forming with the tertiary
veins a prominulous reticulation on the lower side, quite obsolete on
the upper one. Flowers and pedicels cupreous-pilose, fasciculate (5-
12) in the axils of the leaves, more frequently above the leaf-scars;
pedicels (10) -15 (20) mm. long, slightly enlarged above; sepals 5,
quincuncial, the two inner smaller and thinner, the remainder thick-
coriaceous, triangular-concave, 4x3 mm., quite glabrous inside;
corolla very thick, fleshy, 4 mm. high, cupreous-sericeous outside,
glabrous inside, however, with the border glabrous and white, free
lobes slightly imbricate, ovate, 2 mm. high; stamens 5, reduced to
very small and subulate filaments encircled by the thick staminodes,
sparsely sericeous-pilose; pistil stout, 2-2.5 mm. high, glabrous, style
only 0.5 mm. long, stigma inconspicuous, obscurely 4-5 lobed ; ovary
4-5 celled, furrowed, densely sericeous, ovules attached by a very
short and thin funicle from the middle of their adaxial side. Fruit
unknown.

Type: Wurdack 24.25.

Endemic, up to the time of writing, in Amazonian Peru. Loreto:
Rio Maranon, rain forest at upper end of Pongo de Manseriche,
250 m., Wurdack 2425 (F, G, LE). High rain forest along Rio Mar-
anon near Teniente Pinglo, just above Pongo de Manseriche, 250-
300 m., Wurdack 2121 (P).

HECKMAN

BINDERY INC.

OCT96

. PIMJ» N. MANCHESTER,

UNIVERSITY OF ILLINOIS-URBAN*